Assessment of genetic diversity among Jordanian wild barley (Hordeum spontaneum) genotypes revealed by SSR markersby Y. Shakhatreh, M. Baum, N. Haddad, M. Alrababah, S. Ceccarelli

Genet Resour Crop Evol

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RESEARCH ARTICLE

Assessment of genetic diversity among Jordanian wild barley (Hordeum spontaneum) genotypes revealed by SSR markers

Y. Shakhatreh . M. Baum . N. Haddad .

M. Alrababah . S. Ceccarelli

Received: 17 February 2015 / Accepted: 29 June 2015  Springer Science+Business Media Dordrecht 2015

Abstract Wild barley, Hordeum spontaneum is the progenitor of cultivated barley. The center of diversity is in the Fertile Crescent in the Near East, where wild barley grows under a wide range of environmental and climatic conditions. Jordan wild barley is expected to harbor important genes that could be used for the improvement of the cultivated varieties especially in their tolerance to drought. To assess genetic diversity and allelic variation of Jordanian wild barley, 103 wild barley genotypes collected from different parts of

Jordan along with 29 cultivated barley genotypes were analyzed for diversity by means of Simple Sequence

Repeats (SSR). The spontaneum genotypes were grouped into six populations according to their longitude, latitude, altitude, and rainfall zone of the sites from where they have been collected, and the cultivated in one population. All barley genotypes were analyzed with 11 SSR markers with known sequences and chromosomal locations. The 11 microsatellite markers revealed 237 alleles, with an average of 21.5 alleles per locus. In spontaneum genotypes, 209 alleles were identified with an average of 19 alleles per locus, whereas, cultivated genotypes had 95 alleles with an average of 8.6 alleles per locus.

Moreover, 52 alleles were identified in spontaneum (22 %) of the total alleles and 22 alleles in cultivated barley (9 %). The level of genetic diversity was very high; in fact, mean in total gene diversity (Ht) was 0.86 ranging from 0.72 to 0.94. Average gene diversity (H) was 0.79 ranging from 0.74 to 0.82. Genetic variation within population was much higher than among populations at molecular levels: this was also supported by the finding that morphological characters such as awn length, plant height, days to maturity, peduncle length, peduncle extrusion and tillering number, showed higher variation within populations than among populations. Clustering of populations was according to their ecological geographical pattern.

Keywords Drought stress  Genetic diversity 

Hordeum spontaneum  Microsatellite markers 

Wild barley

Introduction

Wild barley, (Hordeum spontaneum C. Koch, syn.:

Hordeum vulgare L. subsp. spontaneum (C. Koch)

Y. Shakhatreh (&)

National Center for Agricultural Research and Extension (NCARE), Amman, Jordan e-mail: shakhatreh12@yahoo.com

M. Baum  N. Haddad

International Center for Agricultural Research in the Dry

Areas (ICARDA), Amman, Jordan

M. Alrababah

Faculty of Agriculture, Jordan University of Science and

Technology (JUST), Irbid, Jordan

S. Ceccarelli

Via delle Begonie 2, 63100 Ascoli Piceno, Italy 123

Genet Resour Crop Evol

DOI 10.1007/s10722-015-0285-8

Thell.) is the progenitor of domesticated barley (Harlan and Zohary 1966; von Bothmer et al. 2003). It occupies diverse habitats ranging from high rainfall regions to deserts (Volis et al. 2002). Barley domestication began in the Fertile Crescent, which includes parts of Iran,

Turkey, Syria, Palestine, and Jordan (Faccioli et al. 1998; Ivandic et al. 2002; von Bothmer et al. 2003).

Wild and cultivated barley are morphologically similar; with the latter, having broader leaves, shorter stem and awns, tough ear rachis, shorter and thicker spike, and larger grains. H. spontaneum is a valuable source of new genes for breeding. These include, grain protein content (Jaradat 1991), resistance to powdery mildew (Dreiseitl and Dinoor 2004), earliness (Korff et al. 2004;Li et al. 2005), biomass (Lu et al. 1999), plant height under drought (Baum et al. 2003), drought tolerance (Baum et al. 2003; Korff et al. 2004; Hu¨bner et al. 2009; Shakhatreh et al. 2010; Lakew et al. 2013), cold tolerance (Grossi et al. 1998; Crosatti et al. 1996), deep-seeding tolerance and fast emergence of seedlings from deep seeded conditions (Takahashi et al. 2001).

The level of genetic variation inH. spontaneum has been studied and estimated using phenotypic characters, isozymes and different PCR-based molecular markers (Turpeinen et al. 2003; Jana and Pietrzak 1988; Dreiseitl and Dinoor 2004; Jaradat 1991; Baum et al. 2004; and Bedada et al. 2014). These studies concluded that the genetic variation in wild barley is substantial.

One of the main objectives in barley improvement is to increase abiotic stress tolerance to enhance crop reliability (Ellis et al. 2000) especially in a time of climate change in the dry areas. Wild barley could be an important source for genes to achieve breeders objectives. One of the first steps in utilizing wild barley is the evaluation of genetic diversity.

The collection from the Fertile Crescent, from

Jordan and the neighboring countries, contains high levels of genetic variation and therefore is an obvious choice for studies related to genetic variation assessment. This is true because Jordan is considered as a center of diversity and origin as indicated by Baek et al. (2003). ‘‘Only Jordan and Syria have high genetic diversity of wild barley similar to Israel’’. As for morphological traits, plant height, biomass yield, earliness, peduncle length, peduncle extrusion and awn length are among the important morphological traits to increase abiotic stress tolerance (Ellis et al. 2000 and Baum et al. 2003; Shakhatreh et al. 2010).

There are several molecular tools for estimating genetic variation, however, Simple Sequence Repeat (SSR) markers are ideal genetic markers for detecting differences between and within species. Microsatellites, SSRs, are short (2–5 nucleotides) tandemly repeated DNA sequences. They occur frequently and almost randomly in most eukaryotic genomes (Tatuz and Renz 1984). Microsatellites can be found anywhere in the genome, both in protein-coding and noncoding regions and even in promoter regions. Since the flanking sequences of each SSR locus are usually unique, primers targeting these flanking regions can be designed to produce an SSR marker (Beckmann and