Skip to main content
    • Aa
    • Aa

Changes in barley (Hordeum vulgare L. subsp. vulgare) genetic diversity and structure in Jordan over a period of 31 years

  • I. Thormann (a1), P. Reeves (a2), S. Thumm (a3), A. Reilley (a2), J. M. M. Engels (a1), C. M. Biradar (a4), U. Lohwasser (a3), A. Börner (a3), K. Pillen (a5) and C. M. Richards (a2)...

In many regions of the world, the cultivation of landraces is still common, in particular in centres of crop diversity. Significant effort has been put into ex situ conservation of landraces but limited data exist on the changes in genetic diversity that occur over time in farmers’ fields. We assessed temporal changes in barley landrace diversity in Jordan using seed samples collected in 1981 and 2012 from the same locations. We did not observe significant changes in the amount of genetic diversity, but samples collected in 2012 were more homogenous and less locally distinct. In two sites, we observed replacement of the old material. We observed a change in phenotype, and phenotypes were found to be more homogeneous among sites in 2012. Climate changed significantly over the study period, becoming hotter and dryer, but we did not identify any correlation between the changes in climate and genetic and phenotypic variations. While the amount of genetic diversity in terms of allelic richness and number of multi-locus genotypes has been maintained, local distinctiveness among landrace barley populations in Jordan was reduced.

Corresponding author
* Corresponding author. E-mail:
Hide All
AbebeTD and LeonJ (2013) Spatial and temporal genetic analyses of Ethiopian barley (Hordeum vulgare L.) landraces reveal the absence of a distinct population structure. Genetic Resources and Crop Evolution 60: 15471558.
AbebeTD, MathewB and LéonJ (2013) Barrier analysis detected genetic discontinuity among Ethiopian barley (Hordeum vulgare L.) landraces due to landscape and human mobility on gene flow. Genetic Resources and Crop Evolution 60: 297309.
Al-BakriJT, TaylorJC and BrewerTR (2001) Monitoring land use change in the Badia transition zone in Jordan using aerial photography and satellite imagery. The Geographical Journal 167: 248262.
Al-BakriJT, AjlouniM and Abu-ZanatM (2008) Incorporating land use mapping and participation in Jordan. Mountain Research and Development 28: 4957.
AlerciaA, DiulgheroffS and MackayM (2015) FAO/Bioversity Multi-Crop Passport Descriptors. Rome, Italy: Food and Agriculture Organization of the United Nations, Bioversity International.
AllinneC, MariacC, VigourouxY, BezançonG, CouturonE, MoussaD, TidjaniM, PhamJ-L and RobertT (2008) Role of seed flow on the pattern and dynamics of pearl millet (Pennisetum glaucum [L.] R. Br.) genetic diversity assessed by AFLP markers: a study in south-western Niger. Genetica 133: 167178.
Al-TabbalJA and Al-FraihatAH (2012) Genetic variation, heritability, phenotypic and genotypic correlation studies for yield and yield components in promising barley genotypes. Journal of Agricultural Science 4: 193201.
Al-YassinA (2012) Jordan: in search of new benefit-sharing practices through participatory plant breeding. In: RuizM and VernooyR (eds) The Custodians of Biodiversity. Sharing Access and Benefits to Genetic Resources. Abingdon, UK: Earthscan, pp. 6778.
AssefaA and LabuschagneMT (2004) Phenotypic variation in barley (Hordeum vulgare L.) landraces from north Shewa in Ethiopia. Biodiversity and Conservation 13: 14411451.
BarryMB, PhamJL, BéavoguiS, GhesquièreA and AhmadiN (2008) Diachronic (1979–2003) analysis of rice genetic diversity in Guinea did not reveal genetic erosion. Genetic Resources and Crop Evolution 55: 723733.
BekeleE (1983) Some measures of gene diversity analysis on land race populations of Ethiopian barley. Hereditas 98: 127143.
BertoldoJG, Meirelles CoimbraJL, GuidolinAF, Braatz de AndradeLR and NodariRO (2014) Agronomic potential of genebank landrace elite accessions for common bean. Sci. Agric. 71: 120125.
BezançonG, PhamJL, DeuM, VigourouxY, SagnardF, MariacC, KapranI, MamadouA, GérardB, NdjeungaJ and ChantereauJ (2009) Changes in the diversity and geographic distribution of cultivated millet (Pennisetum glaucum (L.) R. Br.) and sorghum (Sorghum bicolor (L.) Moench) varieties in Niger between 1976 and 2003. Genetic Resources and Crop Evolution 56: 223236.
BrownAHD (1999) The genetic structure of crop landraces and the challenge to conserve them in situ on farms. In: BrushSB (ed.) Genes in the Field: On-Farm Conservation of Crop Diversity. Ottawa, Canada: IDRC, pp. 2948.
BrownAHD and MarshallDR (1995) A basic sampling strategy: theory and practice. In GuarinoL, Ramanat ha RaoV and ReidR (eds) Collecting Plant Genetic Diversity: Technical Guidelines. Rome, Italy: International Plant Genetic Resources Institute (IPGRI), Plant Production and Protection Division. Rome, Italy: FAO, World Conservation Union (IUCN). Gland, Switzerland: CABI Publishing, Wallingford, UK, pp 7591.
BrownAHD and MundayJ (1982) Population-genetic structure and optimal sampling of land races of barley from Iran. Genetica 58: 8596.
BrushSB (2004) Farmers’ Bounty: Locating Crop Diversity in the Contemporary World. New Haven, USA: Yale University Press.
Camacho VillaTC, MaxtedN, ScholtenM and Ford-LloydB (2005) Defining and identifying crop landraces. Plant Genetic Resources: Characterization and Utilization 3: 373384.
CeccarelliS (1996) Adaptation to low/high input cultivation. Euphytica 92: 203214.
CeccarelliS and GrandoS (1999) Barley landraces from the Fertile Crescent: a lesson for plant breeders. In: BrushSB (ed.) Genes in the Field: On-Farm Conservation of Crop Diversity. Ottawa, Canada: IDRC, pp. 5176.
CeccarelliS and GrandoS (2007) Decentralized-participatory plant breeding: an example of demand driven research. Euphytica 155: 349360.
CeccarelliS and GrandoS (2009) Participatory plant breeding in cereals. In: CarenaMJ (ed.) Cereals. US: Springer, pp. 395414.
CeccarelliS, GrandoS and van LeurJAG (1987) Genetic diversity in barley landraces from Syria and Jordan. Euphytica 36: 389405.
CuiD, LiJ, TangC, AX, YuT, MaX, ZhangE, CaoG, XuF, QiaoY, DaiL and HanL (2016) Diachronic analysis of genetic diversity in rice landraces under on-farm conservation in Yunnan, China. Theoretical and Applied Genetics 129: 155168.
DamaniaAB, JackksonMT and PorcedduE (1985) Variation in wheat and barley landraces from Nepal and the Yemen Arab Republic. Z. Pflanzenzüchtungg 94: 1324.
DavisMB, ShawRG and EttersonJR (2005) Evolutionary responses to changing climate. Ecology 86: 17041714.
Del RioAH, BambergJB, HuamanZ, SalasA and VegaSE (1997) Assessing changes in the genetic diversity of potato genebanks. 2. In situ vs ex situ. Theoretical and Applied Genetics 95: 199204.
DemissieA and BjornstadA (1996) Phenotypic diversity of Ethiopian barleys in relation to geographic regions, altitudinal range, and agro-ecological zones: as an aid to germplasm collection and conservation strategy. Hereditas 124: 1729.
DeuM, SagnardF, ChantereauJ, CalatayudC, VigourouxY, PhamJL, MariacC, KapranI, MamadouA, GérardB, NdjeungaJ and BezançonG (2010) Spatio–temporal dynamics of genetic diversity in Sorghum bicolor in Niger. Theoretical and Applied Genetics 120: 13011313.
DornelasM, MagurranAE, BucklandST, ChaoA, ChazdonRL, ColwellRK, CurtisT, GastonKJ, GotelliNJ, KosnikMA, McGillB, McCuneJL, MorlonH, MumbyPJ, ØvreasL, StudenyA and VellendM (2013) Quantifying temporal change in biodiversity: challenges and opportunities. Proceedings of the Royal Society B 280: 20121931.
DullooME, HansonJ, JorgeMA and ThormannI (2008) Regeneration guidelines: general guiding principles. In: DullooME, ThormannI, JorgeMA and HansonJ (eds) Crop Specific Regeneration Guidelines [CD-ROM]. Rome, Italy: CGIAR System-wide Genetic Resource Programme (SGRP), 6 pp.
DwivediSL, CeccarelliS, BlairMW, UpadhyayaHD, AreAK and OrtizR (2016) Landrace germplasm for improving yield and abiotic stress adaptation. Trends in Plant Science 21: 3142.
ErstsPJ [Internet] Geographic Distance Matrix Generator (version 1.2.3). American Museum of Natural History, Center for Biodiversity and Conservation. Available at Accessed 4 February 2016.
EvannoG, RegnautS and GoudetJ (2005) Detecting the number of clusters of individuals using the software STRUCTURE: a simulation study. Molecular Ecology 14: 26112620.
FAO (2006) Country Pasture/Forage Resource Profiles. Rome, Jordan, Italy: Food and Agriculture Organization of the United Nations.
FAO (2010) The Second Report on the State of the World's Plant Genetic Re-Sources for Food and Agriculture. Rome, Italy: Food and Agriculture Organization of the United Nations.
FAO (2012) Second Global Plan of Action for Plant Genetic Resources for Food and Agriculture. Rome, Italy: Food and Agriculture Organization of the United Nations.
FranksSJ, AviseJC, BradshawWE, ConnerJK, EttersonJR, MazerSJ, ShawRG and WeisAE (2008) The Resurrection Initiative: storing ancestral genotypes to capture evolution in action. BioScience 58: 870873.
FriedtW, HorsleyRD, HarveyBL, PoulsenDME, LanceRCM, CeccarelliS, GrandoS and CapettiniF (2011) Barley breeding history, progress, objectives and technology. In: UllrichSE (ed.) Barley: Production, Improvement and Uses. Ames, USA: Blackwell Publishing Ltd., pp. 160220.
GaoH, WilliamsonS and BustamanteCD (2007) A Markov chain Monte Carlo approach for joint inference of population structure and inbreeding rates from multilocus genotype data. Genetics 176: 16351651.
GomezOJ, BlairMW, Frankow-LindbergBE and GullbergU (2005) Comparative study of common bean (Phaseolus vulgaris L.) landraces conserved ex situ in genebanks and in situ by farmers. Genetic Resources and Cop Evolution 52: 371380.
GoudetJ (2001) FSTAT, a program to estimate and test gene diversities and fixation indices (Version 2.9.3). Available at
GuoY, LiY, HuangY, JarvisD, SatoK, KatoK, TsuyuzakiH, ChenL and LongC (2012) Genetic diversity analysis of hulless barley from Shangri-la region revealed by SSR and AFLP markers. Genetic Resources and Crop Evolution 59: 15431552.
HaseneyerG, StrackeS, PaulC, EinfeldtC, BrodaA, PiephoH-P, GranerA and GeigerHH (2010) Population structure and phenotypic variation of a spring barley world collection set up for association studies. Plant Breeding 129: 271279.
HenryJP, PontisC, DavidJ and GouyonPH (1991) An experiment on dynamic conservation of genetic resources with metapopulations. In: SeitzA, LoeschckeV (eds) Species Conservation: A Population-Biological Approach. Basel, Switzerland: Birkhäuser Verlag Basel, pp. 185198.
HobanS and SchlarbaumS (2014) Optimal sampling of seeds from plant populations for ex-situ conservation of genetic biodiversity, considering realistic population structure. Biological Conservation 177: 9099.
HobanS, ArntzenJA, BrufordMW, GodoyJA, HoelzelR, SegelbacherG, VilaC and BertorelleG (2014) Comparative evaluation of potential indicators and temporal sampling protocols for monitoring genetic erosion. Evolutionary Applications 7: 984998.
HolmS (1979) A simple sequentially rejective multiple test procedure. Scandinavian Journal of Statistics 6: 6570.
ICARDA (2003) From formal to participatory plant breeding: improving barley production in the rainfed areas of Jordan. Final technical report. International Center for Agricultural Research in the Dry Areas. Available at accessed 22 April 2016.
JakobssonM and RosenbergNA (2007) CLUMPP: a cluster matching and permutation program for dealing with label switching and multimodality in analysis of population structure. Bioinformatics 23: 18011806.
JanaS and PietrzakLN (1988) Comparative assessment of genetic diversity in wild and primitive cultivated barley in a center of diversity. Genetics 119: 981990.
JaradatAA (1989a) Diversity within and between populations of two sympatrically distributed Hordeum species in Jordan. Theoretical and Applied Genetics 78: 653656.
JaradatAA (1989b) Ecotypes and genetic divergence among sympatrically distributed populations of Hordeum vulgare and Hordeum spontaneum from the xeric region of Jordan. Theoretical and Applied Genetics 78: 857862.
JarvisDI, MyerL, KlemickH, GuarinoL, SmaleM, BrownAHD, SadikiM, SthapitB and HodgkinT (2000) A Training Guide for In Situ Conservation On-farm. Version 1. Rome, Italy: International Plant Genetic Resources Institute.
JarvisDI, HodgkinT, SthapitBR, FaddaC and Lopez-NoriegaI (2011) An heuristic framework for identifying multiple ways of supporting the conservation and use of traditional crop varieties within the agricultural production system. Critical Reviews in Plant Sciences 30: 125176.
JostL (2008) G ST and its relatives do not measure differentiation. Molecular Ecology 17: 40154026.
KeenanK, McGinnityP, CrossTF, CrozierWW and ProdöhlPA (2013) diveRsity: an R package for the estimation of population genetics parameters and their associated errors. Methods in Ecology and Evolution 4: 782788.
KhlestkinaEK, HuangXQ, QuenumFJ-B, ChebotarS, RoederMS and BörnerA (2004) Genetic diversity in cultivated plants – loss or stability? Theoretical and Applied Genetics 108: 14661472.
KhlestkinaEK, VarshneyRK, RöderMS, GranerA and BörnerA (2006) A comparative assessment of genetic diversity in cultivated barley collected in different decades of the last century in Austria, Albania and India by using genomic and genic simple sequence repeat (SSR) markers. Plant Genetic Resources: Characterization and Utilization 4: 125133.
KhresatSA, RawajfihZ and MohammadM (1998) Land degradation in north-western Jordan: causes and processes. Journal of Arid Environments 39: 623629.
KirkpatrickM and LandeR (1989) The evolution of maternal characters. Evolution 43: 485503.
LasaJM, IgartuaE, CiudadFJ, CodesalP, GarcíaEV, GraciaMP, MedinaB, RomagosaI, Molina-CanoJL and MontoyaJL (2001) Morphological and agronomical diversity patterns in the Spanish barley core collection. Hereditas 135: 217225.
LewisPO and ZaykinD (2001) Genetic Data Analysis: Computer program for the analysis of allelic data (version 1.1). Available at
Malysheva-OttoLV, GanalMW and RöderMS (2006) Analysis of molecular diversity, population structure and linkage disequilibrium in a worldwide survey of cultivated barley germplasm (Hordeum vulgare L.). BMC Genetics 7: 6. DOI: 10.1186/1471-2156-7-6.
MaxtedM and GuarinoL (2006) Genetic erosion and genetic pollution of crop wild relatives. In: Ford-LloydBV, DiasSR and BettencourtE (eds) Genetic Erosion and Pollution Assessment Methodologies. Proceedings of PGR Forum Workshop 5, Terceira Island, Autonomous Region of the Azores, Portugal, 8–11 September 2004. Rome: Published on behalf of the European Crop Wild Relative Diversity Assessment and Conservation Forum, Bioversity International, pp. 3546.
MegersaG (2014) Genetic erosion of barley in North Shewa Zone of Oromiya Region, Ethiopia. International Journal of Biodiversity and Conservation 6: 280289.
MeirmansPG and HedrickPW (2011) Assessing population structure: FST and related measures. Molecular Ecology 11: 518.
MekbibF (2008) Genetic erosion of sorghum (Sorghum bicolor (L.) Moench) in the centre of diversity, Ethiopia. Genetic Resources and Crop Evolution 55: 351364.
MercerKL and PeralesHR (2010) Evolutionary response of landraces to climate change in centers of crop diversity. Evolutionary Applications 3: 480493.
MousseauTA and FoxCW (1998) The adaptive significance of maternal effects. Trends in Ecology & Evolution 13: 403407.
NagelM, VogelH, LandjevaS, Buck-SorlinG, LohwasserU, ScholzU and BörnerA (2009) Seed conservation in ex-situ genebanks - genetic studies on longevity in barley. Euphytica 170: 110.
NCARTT (2007) The Second Country Report on the State of the Plant Genetic Resources for Food and Agriculture. The Hashemite Kingdom of Jordan. Amman, Jordan: National Center for Agricultural Research and Technology Transfer NCARTT.
Ould Med MahmoudMA and HamzaS (2009) Genetic diversity in local barley accessions collected from different geographical regions of Tunisia. Plant Genetic Resources: Characterization and Utilization 7: 169176.
PandeyM, WagnerC, FriedtW and OrdonF (2006) Genetic relatedness and population differentiation of Himalayan hulless barley (Hordeum vulgare L.) landraces inferred with SSRs. Theoretical and Applied Genetics 113: 715729.
ParziesHK, SpoorW, EnnosAR (2000) Outcrossing rates of barley landraces from Syria. Plant Breeding 119: 520522.
ParziesHK, SpoorW and EnnosAR (2004) Inferring seed exchange between farmers from population genetic structure of barley landrace Arabi Aswad from Northern Syria. Genetic Resources and Crop Evolution 51: 471478.
PeakallR and SmousePE (2006) GENALEX 6: genetic analysis in Excel. Population genetic software for teaching and research. Molecular Ecology Notes 6: 288295.
PeakallR and SmousePE (2012) GenAlEx 6.5: genetic analysis in Excel. Population genetic software for teaching and research-an update. Bioinformatics 28: 25372539.
Pérez de la VegaM and GarciaP (1997) Genetic structure of self-pollinating species: the case of wild Avena . Bocconea 7: 141152.
PetrovicS and DimitrijevicM (2012) Genetic erosion of diversity in cereals. Genetika 44: 217226.
PritchardJK, StephensM and DonnellyP (2000) Inference of population structure using multilocus genotype data. Genetics 155: 945959.
RaoNK, HansonJ, DullooME, GhoshK, NowellD and LarindeM (2006) Manual of Seed Handling in Genebanks. Handbooks for Genebanks No. 8. Rome, Italy: Bioversity International.
RosenbergNA (2004) Distruct: a program for the graphical display of population structure. Molecular Ecology Notes 4: 137138.
RoussetF (1997) Genetic differentiation and estimation of gene flow from F-statistics under isolation by distance. Genetics 145: 12191228.
RussellJR, BoothA, FullerJD, BaumM, CeccarelliS, GrandoS and PowellW (2003) Patterns of polymorphism detected in the chloroplast and nuclear genomes of barley landraces sampled from Syria and Jordan. Theoretical and Applied Genetics 107: 413421.
RussellJ, DawsonIK, FlavellAJ, SteffensonB, WeltzienE, BoothA, CeccarelliS, GrandoS and WaughR (2011) Analysis of 1000 single nucleotide polymorphisms in geographically matched samples of landrace and wild barley indicates secondary contact and chromosome-level differences in diversity around domestication genes. New Phytologist 191: 564578.
SambergLH, FishmanL and AllendorfFW (2013) Population genetic structure in a social landscape: barley in a traditional Ethiopian agricultural system. Evolutionary Applications 6: 11331145.
SteeleKA, GyawaliS, JoshiKD, ShresthaP, SthapitBR and WitcombeJR (2009) Has the introduction of modern rice varieties changed rice genetic diversity in a high-altitude region of Nepal?. Field Crops Research 113: 2430.
SteinN, PrasadM, ScholzU, ThielT, ZhangH, WolfM, KotaR, VarshneyRK, PerovicD, GrosseI and GranerA (2007) A 1000-loci transcript map of the barley genome: new anchoring points for integrative grass genomics. Theoretical and Applied Genetics 114: 823839.
ThielT, MichalekW, VarshneyRK and GranerA (2003) Exploiting EST databases for the development and characterization of gene-derived SSR-markers in barley (Hordeum vulgare L.). Theoretical and Applied Genetics 106: 411422.
ThomasM, DawsonJC, GoldringerI and BonneuilC (2011) Seed exchanges, a key to analyze crop diversity dynamics in farmer-led on-farm conservation. Genetic Resources and Crop Evolution 58: 321338.
ThormannI and EngelsJMM (2015) Genetic diversity and erosion – a global perspective. Chapter 10. In: AhujaMR and JainSM (eds) Genetic Diversity and Erosion – Indicators and Prevention, vol. 1. Switzerland: Springer International Publishing, pp. 263294.
ThormannI, FiorinoE, HalewoodM and EngelsJMM (2015) Plant genetic resources collections and associated information as baseline resource for genetic diversity studies – an assessment of the IBPGR supported collections. Genetic Resources and Crop Evolution 62: 12791293.
Van de WouwM, KikC, van HintumT, van TreurenR and VisserB (2010) Genetic erosion in crops: concept, research, results and challenges. Plant Genetic Resources: Characterization and Utilization 8: 115.
Van HintumT and MentingF (2003) Diversity in ex situ genebank collections of barley. In: von BothmerR, van HintumT and KnüpfferH (eds) Diversity in Barley (Hordeum vulgare). Developments in Plant Genetics and Breeding, vol. 7. Amsterdam: Elsevier, pp. 247257.
VarshneyRK, MarcelTC, RamsayL, RussellJ, RöderM, SteinN, WaughR, LangridgeP, NiksRE and GranerA (2007) A high density barley microsatellite map with 775 SSR loci. Theoretical and Applied Genetics 114: 10911103.
VarshneyRK, SalemKFM, BaumM, RöderMS, GranerA and BörnerA (2008) SSR and SNP diversity in a barley germplasm collection. Plant Genetic Resources: Characterization and Utilization 6: 167174.
VigourouxY, MariacC, De MitaS, PhamJ-L, GérardB, KapranI, SagnardF, DeuM, ChantereauJ, AliA, NdjeungaJ, LuongV, ThuilletAC, SaidouAA and BezanconG (2011) Selection for Earlier Flowering Crop Associated with Climatic Variations in the Sahel. PLoS ONE 6: e19563.
WagnerDB and AllardRW (1991) Pollen migration in predominantly self-fertilizing plants: barley. Journal of Heredity 82: 302304.
WeltzienE (1982) Barley collection and evaluation in Syria and Jordan. Plant Genetic Resources Newsletter 52: 56.
WeltzienE (1988) Evaluation of barley (Hordeum vulgare L.) landrace populations originating from different growing regions in the Near East. Plant Breeding 101: 95106.
WestengenOT, OkongoMA, OnekL, BergT, Hai UpadhyayaH, BirkelandS, KhalsaSDK, RingKH, NilsC, StensethNC and BrystingAK (2014) Ethnolinguistic structuring of sorghum genetic diversity in Africa and the role of local seed systems. PNAS 111: 1410014105.
WitcombeJR, BourgoisJJ and RifaieR (1982) Germplasm collections from Syria & Jordan. Plant Genetic Resources Newsletter 50: 28.
YahiaouiS, IgartuaE, MoralejoM, RamsayL, Molina-CanoJL, CiudadFJ, LasaJM, GraciaMP and CasasAM (2008) Patterns of genetic and eco-geographical diversity in Spanish barleys. Theoretical and Applied Genetics 116: 271282.
Recommend this journal

Email your librarian or administrator to recommend adding this journal to your organisation's collection.

Plant Genetic Resources
  • ISSN: 1479-2621
  • EISSN: 1479-263X
  • URL: /core/journals/plant-genetic-resources
Please enter your name
Please enter a valid email address
Who would you like to send this to? *


Type Description Title
Supplementary Materials

Thormann supplementary material
Thormann supplementary material

 Unknown (251 KB)
251 KB


Full text views

Total number of HTML views: 6
Total number of PDF views: 42 *
Loading metrics...

Abstract views

Total abstract views: 507 *
Loading metrics...

* Views captured on Cambridge Core between 15th May 2017 - 22nd October 2017. This data will be updated every 24 hours.