In this study, simple sequence repeat (SSR) markers were used to investigate the genetic diversity and evolutionary relationships among 74 barley accessions; 15 landraces (Hordeum vulgare) and 59 wild accessions (twenty-one Hordeum spontaneum, ten Hordeum bulbosum, thirteen Hordeum murinum and fifteen Hordeum marinum). A total of 66 alleles were detected at 12 SSR loci, with an average of 5.5 alleles per locus in the entire samples. The average number of alleles (4.2) per locus was higher in H. vulgare and H. spontaneum, reflecting more genetic diversity within these two species compared with the other species. All index values of genetic diversity revealed that there was higher genetic diversity within H. vulgare landraces than within the wild species, indicating that cultivation unlikely to have caused a loss of genetic diversity in cultivated barley compared with its wild relatives. Analysis of molecular variance (AMOVA) revealed that 31% of the total observed variation was accounted for among species. Based on neighbor-joining clustering, the five species were classified into three main groups: (i) all accessions of H. marinum and H. bulbosum plus two accessions of H. murinum; (ii) H. vulgare and H. spontaneum accessions, in small, separated groups; and (iii) the remaining accessions of H. murinum. Principal coordinates analysis (PCoA) for SSR data also supported the neighbor-joining clustering. These results have important implications for barley germplasm characterization, improvement, molecular systematics and conservation.
Baek, H.J., Beharav, A., Nevo, E. 2003. Ecological-genomic diversity of microsatellites in wild barley, Hordeum spontaneum populations in Jordan. Theor. Appl. Genet. 106:397–410.
Nevo E. , 'Ecological-genomic diversity of microsatellites in wild barley, Hordeum spontaneum populations in Jordan ' (2003 ) 106 Theor. Appl. Genet. : 397 -410.
Blattner, F.R. 2004. Phylogenetic analysis of Hordeum (Poaceae) as inferred by nuclear rDNA ITS sequences. Mol. Phylogenet. Evol. 33:289–299.
Blattner F.R. , 'Phylogenetic analysis of Hordeum (Poaceae) as inferred by nuclear rDNA ITS sequences ' (2004 ) 33 Mol. Phylogenet. Evol. : 289 -299.
Dreiseitl, A., Bockelman, H.E. 2004. Sources of powdery mildew resistance in a wild barley collection. Genet. Resour. Crop Evol. 50:345–350.
Bockelman H.E. , 'Sources of powdery mildew resistance in a wild barley collection ' (2004 ) 50 Genet. Resour. Crop Evol. : 345 -350.
Ellis, R.P., Forster, B.P., Robinson, D., Handley, L.L., Gordon, D.C., Russell, J.R., Powell, W. 2000. Wild barley: A source of genes for crop improvement in the 21st century? J. Exp. Bot. 51:9–17.
Powell W. , 'Wild barley: A source of genes for crop improvement in the 21st century? ' (2000 ) 51 J. Exp. Bot. : 9 -17.
Feuerstein, U., Brown, A.H.D., Burdon, J.J. 2006. Linkage of rust resistance genes from wild barley (Hordeum spontaneum) with isozyme markers. Plant Breeding 104:318–324.
Burdon J.J. , 'Linkage of rust resistance genes from wild barley (Hordeum spontaneum) with isozyme markers ' (2006 ) 104 Plant Breeding : 318 -324.
Garthwaite, A.J., von Bothmer, R., Colmer, T.D. 2005. Salt tolerance in wild Hordeum species is associated with restricted entry of Na+ and Cl− into the shoots. J. Exp. Bot. 56:2365–2378.
Colmer T.D. , 'Salt tolerance in wild Hordeum species is associated with restricted entry of Na+ and Cl− into the shoots ' (2005 ) 56 J. Exp. Bot. : 2365 -2378.
Hamza, S., Hamida, W.B., Rebaï, A., Harrabi, M. 2004. SSR-based genetic diversity assessment among Tunisian winter barley and relationship with morphological traits. Euphytica 135:107–118.
Harrabi M. , 'SSR-based genetic diversity assessment among Tunisian winter barley and relationship with morphological traits ' (2004 ) 135 Euphytica : 107 -118.
Harlan, J.R. 1968. On the origin of barley. In: USDA Agriculture Handbook 338:9–31.
Harlan J.R. , 'On the origin of barley ' (1968 ) 338 USDA Agriculture Handbook : 9 -31.
Harlan, J.R. 1995. Barley. In: Smartt, J., Simmonds, N.W. (eds), Evolution of Crop Plants, 2nd ed., Longman, London, UK, pp. 140–146.
Jana, S., Pietrzak, L.N. 1988. Comparative assessment of genetic diversity in wild and primitive cultivated barley in a centre of diversity. Genetics 119:981–990.
Pietrzak L.N. , 'Comparative assessment of genetic diversity in wild and primitive cultivated barley in a centre of diversity ' (1988 ) 119 Genetics : 981 -990.
Jilal, A., Grando, S., Henry, R.J., Lee, L.S., Rice, N., Hill, H., Baum, M., Ceccarelli, S. 2008. Genetic diversity of ICARDA’s worldwide barley landrace collection. Genet. Resour. Crop Evol. 55:1221–1230.
Ceccarelli S. , 'Genetic diversity of ICARDA’s worldwide barley landrace collection ' (2008 ) 55 Genet. Resour. Crop Evol. : 1221 -1230.
Liu, Z.W., Biyashev, R.M., Saghai-Maroof, M.A. 1996. Development of simple sequence repeat DNA markers and their integration into a barley linkage map. Theor. Appl. Genet. 93:869–876.
Saghai-Maroof M.A. , 'Development of simple sequence repeat DNA markers and their integration into a barley linkage map ' (1996 ) 93 Theor. Appl. Genet. : 869 -876.
Manjunatha, T., Bisht, I.S., Bhat, K.V., Singh, B.P. 2007. Genetic diversity in barley (Hordeum vulgare L. ssp. vulgare) landraces from Uttaranchal Himalaya of India. Genet. Resour. Crop Evol. 54:55–65.
Singh B.P. , 'Genetic diversity in barley (Hordeum vulgare L. ssp. vulgare) landraces from Uttaranchal Himalaya of India ' (2007 ) 54 Genet. Resour. Crop Evol. : 55 -65.
Marmiroli, N., Maestri, E., Liviero, L., Massari, A., Malcevschi, A., Monciardini, P. 1999. Application of genomics in assessing biodiversity in wild and cultivated barley. Mol. Ecol. 8:95–106.
Monciardini P. , 'Application of genomics in assessing biodiversity in wild and cultivated barley ' (1999 ) 8 Mol. Ecol. : 95 -106.
Matus, I.A., Hayes, P.M. 2002. Genetic diversity in three groups of barley germplasm assessed by simple sequence repeats. Genome 45:1095–1106.
Hayes P.M. , 'Genetic diversity in three groups of barley germplasm assessed by simple sequence repeats ' (2002 ) 45 Genome : 1095 -1106.
Morrell, P.L., Clegg, M.T. 2007. Genetic evidence for a second domestication of barley (Hordeum vulgare) east of the Fertile Crescent. Proc. Natl. Acad. Sci. USA 104:3289–3294.
Clegg M.T. , 'Genetic evidence for a second domestication of barley (Hordeum vulgare) east of the Fertile Crescent ' (2007 ) 104 Proc. Natl. Acad. Sci. USA : 3289 -3294.
Naghavi, M.R., Aghaei, M.J., Taleei, A.R., Omidi, M., Mozafari, J., Hassani, M.E. 2009. Genetic diversity of the D-genome in T. aestivum and Aegilops species using SSR markers. Genet. Resour. Crop Evol. 56:499–506.
Hassani M.E. , 'Genetic diversity of the D-genome in T. aestivum and Aegilops species using SSR markers ' (2009 ) 56 Genet. Resour. Crop Evol. : 499 -506.
Neale, D.B., Saghai-Maroof, M.A., Allard, R.W., Zhang, Q., Jorgensen, R.A. 1988. Chloroplast DNA diversity in populations of wild and cultivated barley. Genetics 120:1105–1110.
Jorgensen R.A. , 'Chloroplast DNA diversity in populations of wild and cultivated barley ' (1988 ) 120 Genetics : 1105 -1110.
Nei, M. 1978. Estimation of average heterozygosity and genetic distance from a small number of individuals. Genetics 89:583–590.
Nei M. , 'Estimation of average heterozygosity and genetic distance from a small number of individuals ' (1978 ) 89 Genetics : 583 -590.
Nesbitt, M., Samuel, D. 1996. From staple crop to extinction? The archaeology and history of the hulled wheats. In: Padulosi, D., Hammer, K., Heller, J. (eds), Hulled Wheats (Proceedings of the First International Workshop on Hulled Wheats). International Plant Genetics Resources Institute, Rome, Italy, pp. 41–100.
Samuel D. , '', in Hulled Wheats (Proceedings of the First International Workshop on Hulled Wheats) , (1996 ) -.
Nevo, E. 1992. Origin, evolution, population genetics and resources for breeding of wild barley, Hordeum spontaneum, in the Fertile Crescent. In: Shewry, P.R. (ed.), Barley: Genetics, Biochemistry, Molecular Biology and Biotechnology. C.A.B. International, The Alden Press, Oxford, UK, pp. 19–43.
Nevo E. , '', in Barley: Genetics, Biochemistry, Molecular Biology and Biotechnology , (1992 ) -.
Nevo, E., Beiles, A., Zohary, D. 1986. Genetic resources of wild barley in the Near East: Structure, evolution, and application in breeding. Biol. J. Linn. Soc. 27:355–380.
Zohary D. , 'Genetic resources of wild barley in the Near East: Structure, evolution, and application in breeding ' (1986 ) 27 Biol. J. Linn. Soc. : 355 -380.
Pan, Z.F., Deng, G.B., Zhai, X.G., Long, H., Tang, Y.W., Qiang, X.L., Yu, M.Q. 2008. Molecular analysis of cultivated naked barley (Hordeum vulgare L.) from Qinghai-Tibet Plateau in China using SSR markers. Front. Agric. China 2:372–379.
Yu M.Q. , 'Molecular analysis of cultivated naked barley (Hordeum vulgare L.) from Qinghai-Tibet Plateau in China using SSR markers ' (2008 ) 2 Front. Agric. China : 372 -379.
Pandey, M., Wagner, C., Friedt, W., Ordon, F. 2006. Genetic relatedness and population differentiation of Himalayan hulless barley (Hordeum vulgare L.) landraces inferred with SSRs. Theor. Appl. Genet. 113:715–729.
Ordon F. , 'Genetic relatedness and population differentiation of Himalayan hulless barley (Hordeum vulgare L.) landraces inferred with SSRs ' (2006 ) 113 Theor. Appl. Genet. : 715 -729.
Peakal, R., Smouse, P.E. 2006. GenAlEx 6: Genetic analysis in Excel. Population genetic software for teaching and research. Mol. Ecol. Notes 6:288–295.
Smouse P.E. , 'GenAlEx 6: Genetic analysis in Excel. Population genetic software for teaching and research ' (2006 ) 6 Mol. Ecol. Notes : 288 -295.
Powell, W., Morgante, M., Andre, C., Hanafey, M., Vogel, J., Tingey, S., Rafalski, A. 1996. The utility of RFLP, RAPD, AFLP and SSRP (microsatellite) markers for germplasm analysis? Mol. Breed. 2:225–238.
Rafalski A. , 'The utility of RFLP, RAPD, AFLP and SSRP (microsatellite) markers for germplasm analysis? Mol ' (1996 ) 2 Breed. : 225 -238.
Russell, J., Fuller, J., Young, G., Thomas, W.T.B., Taramino, G., Macaulay, M., Waugh, R., Powell, W. 1997. Discriminating between barley genotypes using microsatellite markers. Genome 40:442–450.
Powell W. , 'Discriminating between barley genotypes using microsatellite markers ' (1997 ) 40 Genome : 442 -450.
Saghai-Maroof, M.A., Soliman, K., Jorgensen, R.A., Allard, R.W. 1984. Ribosomal DNA spacer length polymorphisms in barley: Mendelian inheritance, chromosomal location, and population dynamics. Proc. Natl. Acad. Sci. USA 81:8014–8018.
Allard R.W. , 'Ribosomal DNA spacer length polymorphisms in barley: Mendelian inheritance, chromosomal location, and population dynamics ' (1984 ) 81 Proc. Natl. Acad. Sci. USA : 8014 -8018.
Saghai-Maroof, M.A., Biyashev, R.M., Yang, G.P., Zhang, Q., Allard, R.W. 1994. Extraordinarily polymorphic microsatellite DNA in barley: Species diversity, chromosomal locations and population dynamics. Proc. Natl. Acad. Sci. USA 91:5466–5470.
Allard R.W. , 'Extraordinarily polymorphic microsatellite DNA in barley: Species diversity, chromosomal locations and population dynamics ' (1994 ) 91 Proc. Natl. Acad. Sci. USA : 5466 -5470.
Sonina, V.L., Ushnikovaa, A., Tihonova, P., Ananiev, E.V. 1989. Dialect-I, species-specific repeated DNA sequence from barley, Hordeurn vulgare. Theor. Appl. Genet. 78:589–593.
Ananiev E.V. , 'Dialect-I, species-specific repeated DNA sequence from barley, Hordeurn vulgare ' (1989 ) 78 Theor. Appl. Genet. : 589 -593.
Stephen, J.M., Wheatcroft, R., Fedak, G. 1992. RFLP analysis of Hordeum species relationships. Hereditas 116:87–91.
Fedak G. , 'RFLP analysis of Hordeum species relationships ' (1992 ) 116 Hereditas : 87 -91.
von Bothmer, R. 1992. The wild species of Hordeum: Relationships and potential use for improvement of cultivated barley. In: Shewry, P.R. (ed.), Barley: Genetics, Biochemistry, Molecular Biology and Biotechnology. CAB International, Wallingford, UK, pp. 3–18.
Bothmer R. , '', in Barley: Genetics, Biochemistry, Molecular Biology and Biotechnology , (1992 ) -.
von Bothmer, R., Flink, J., Landstrom, T. 1986. Meiosis in interspecific Hordeum hybrids. I. diploid combinations. Can. J. Genet. Cytol. 28:525–535.
Landstrom T. , 'Meiosis in interspecific Hordeum hybrids. I. diploid combinations ' (1986 ) 28 Can. J. Genet. Cytol. : 525 -535.
von Bothmer, R., Van Hintum, T.J.L., Knupffer, H., Sato, K. 2003. Diversity in Barley (Hordeum vulgare). Elsevier Science, Amsterdam, The Netherlands.
Sato K. , '', in Diversity in Barley (Hordeum vulgare) , (2003 ) -.
Yin, Y.Q., Ma, D.Q., Ding, Y. 2003. Analysis of genetic diversity of hordein in wild close relatives of barley from Tibet. Theor. Appl. Genet. 107:837–842.
Ding Y. , 'Analysis of genetic diversity of hordein in wild close relatives of barley from Tibet ' (2003 ) 107 Theor. Appl. Genet. : 837 -842.
Yu, S.B., Xu, W.J., Vijayakumar, C.H.M., Ali, J., Fu, B.Y., Xu, J.L., Jiang, Y.Z., Marghirang, R., Domingo, J., Aquino, C., Virmani, S.S., Li, Z.K. 2003. Molecular diversity and multilocus organization of the parental lines used in the International Rice Molecular Breeding Program. Theor. Appl. Genet. 108:131–140.
Li Z.K. , 'Molecular diversity and multilocus organization of the parental lines used in the International Rice Molecular Breeding Program ' (2003 ) 108 Theor. Appl. Genet. : 131 -140.
Zhang, Q., Yang, G.P., Dai, X., Sun, J.Z. 1994. A comparative analysis of genetic polymorphism in wild and cultivated barley from Tibet using isozyme and ribosomal DNA markers. Genome 37:631–638.
Sun J.Z. , 'A comparative analysis of genetic polymorphism in wild and cultivated barley from Tibet using isozyme and ribosomal DNA markers ' (1994 ) 37 Genome : 631 -638.
Zohary, D., Hopf, M. 2000. Domestication of Plants in the Old World: The Origin and Spread of Cultivated Plants in West Asia, Europe, and the Nile Valley. Oxford Univ. Press, Oxford, UK, 328 pp.
Hopf M. , '', in Domestication of Plants in the Old World: The Origin and Spread of Cultivated Plants in West Asia, Europe, and the Nile Valley , (2000 ) -.