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Genomic analysis of 6,000-year-old cultivated grain illuminates the domestication history of barley

Nature Genetics volume 48, pages 1089–1093 (2016)Cite this article

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Abstract

The cereal grass barley was domesticated about 10,000 years before the present in the Fertile Crescent and became a founder crop of Neolithic agriculture1. Here we report the genome sequences of five 6,000-year-old barley grains excavated at a cave in the Judean Desert close to the Dead Sea. Comparison to whole-exome sequence data from a diversity panel of present-day barley accessions showed the close affinity of ancient samples to extant landraces from the Southern Levant and Egypt, consistent with a proposed origin of domesticated barley in the Upper Jordan Valley. Our findings suggest that barley landraces grown in present-day Israel have not experienced major lineage turnover over the past six millennia, although there is evidence for gene flow between cultivated and sympatric wild populations. We demonstrate the usefulness of ancient genomes from desiccated archaeobotanical remains in informing research into the origin, early domestication and subsequent migration of crop species.

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Figure 1: Ancient plant remains excavated at Yoram Cave.
Figure 2: Ancient barley samples are closely related to present-day landraces from the Levant.
Figure 3: Relationship between genetic similarity and geographical distance.
Figure 4: Gene flow between wild and domesticated barleys in the Levant.

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References

  1. Zohary, D., Hopf, M. & Weiss, E. Domestication of Plants in the Old World: The Origin and Spread of Domesticated Plants in Southwest Asia, Europe, and the Mediterranean Basin (Oxford University Press on Demand, 2012).

  2. Troy, C.S. et al. Genetic evidence for Near-Eastern origins of European cattle. Nature 410, 1088–1091 (2001).

    Article  CAS  PubMed  Google Scholar 

  3. Larson, G. et al. Ancient DNA, pig domestication, and the spread of the Neolithic into Europe. Proc. Natl. Acad. Sci. USA 104, 15276–15281 (2007).

    Article  PubMed  PubMed Central  Google Scholar 

  4. Skoglund, P., Ersmark, E., Palkopoulou, E. & Dalén, L. Ancient wolf genome reveals an early divergence of domestic dog ancestors and admixture into high-latitude breeds. Curr. Biol. 25, 1515–1519 (2015).

    Article  CAS  PubMed  Google Scholar 

  5. da Fonseca, R.R. et al. The origin and evolution of maize in the Southwestern United States. Nat. Plants 1, 14003 (2015).

    Article  PubMed  Google Scholar 

  6. Harlan, J.R. & Zohary, D. Distribution of wild wheats and barley. Science 153, 1074–1080 (1966).

    Article  CAS  PubMed  Google Scholar 

  7. Poets, A.M., Fang, Z., Clegg, M.T. & Morrell, P.L. Barley landraces are characterized by geographically heterogeneous genomic origins. Genome Biol. 16, 173 (2015).

    Article  PubMed  PubMed Central  Google Scholar 

  8. Badr, A. et al. On the origin and domestication history of barley (Hordeum vulgare). Mol. Biol. Evol. 17, 499–510 (2000).

    Article  CAS  PubMed  Google Scholar 

  9. Li, C. et al. Ancient DNA analysis of desiccated wheat grains excavated from a Bronze Age cemetery in Xinjiang. J. Archaeol. Sci. 38, 115–119 (2011).

    Article  CAS  Google Scholar 

  10. Palmer, S.A., Moore, J.D., Clapham, A.J., Rose, P. & Allaby, R.G. Archaeogenetic evidence of ancient Nubian barley evolution from six to two-row indicates local adaptation. PLoS One 4, e6301 (2009).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  11. Smith, O. et al. Sedimentary DNA from a submerged site reveals wheat in the British Isles 8000 years ago. Science 347, 998–1001 (2015).

    Article  CAS  PubMed  Google Scholar 

  12. Weiß, C.L., Dannemann, M., Prüfer, K. & Burbano, H.A. Contesting the presence of wheat in the British Isles 8,000 years ago by assessing ancient DNA authenticity from low-coverage data. eLife 4, e10005 (2015).

    Article  PubMed  PubMed Central  Google Scholar 

  13. Kistler, L. et al. Thermal age, cytosine deamination and the veracity of 8,000 year old wheat DNA from sediments. Preprint at bioRxiv http://dx.doi.org/10.1101/032060 (2015).

  14. Briggs, A.W. et al. Patterns of damage in genomic DNA sequences from a Neandertal. Proc. Natl. Acad. Sci. USA 104, 14616–14621 (2007).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  15. Sawyer, S., Krause, J., Guschanski, K., Savolainen, V. & Pääbo, S. Temporal patterns of nucleotide misincorporations and DNA fragmentation in ancient DNA. PLoS One 7, e34131 (2012).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  16. Briggs, A.W. et al. Removal of deaminated cytosines and detection of in vivo methylation in ancient DNA. Nucleic Acids Res. 38, e87 (2010).

    Article  PubMed  CAS  Google Scholar 

  17. Mascher, M. et al. Barley whole exome capture: a tool for genomic research in the genus Hordeum and beyond. Plant J. 76, 494–505 (2013).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  18. Russell, J. et al. Exome sequencing of geographically diverse barley landraces and wild relatives gives insights into environmental adaptation. Nat. Genet. http://dx.doi.org/10.1038/ng.3612 (2016).

  19. Patterson, N., Price, A.L. & Reich, D. Population structure and eigenanalysis. PLoS Genet. 2, e190 (2006).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  20. Reich, D., Thangaraj, K., Patterson, N., Price, A.L. & Singh, L. Reconstructing Indian population history. Nature 461, 489–494 (2009).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  21. Pourkheirandish, M. et al. Evolution of the grain dispersal system in barley. Cell 162, 527–539 (2015).

    Article  CAS  PubMed  Google Scholar 

  22. Komatsuda, T. et al. Six-rowed barley originated from a mutation in a homeodomain–leucine zipper I–class homeobox gene. Proc. Natl. Acad. Sci. USA 104, 1424–1429 (2007).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  23. Schiffels, S. et al. Iron Age and Anglo-Saxon genomes from East England reveal British migration history Nat. Commun. 7, 10408 (2016).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  24. Mathieson, I. & McVean, G. Differential confounding of rare and common variants in spatially structured populations. Nat. Genet. 44, 243–246 (2012).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  25. Nordborg, M. et al. The pattern of polymorphism in Arabidopsis thaliana. PLoS Biol. 3, e196 (2005).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  26. Morrell, P.L. & Clegg, M.T. Genetic evidence for a second domestication of barley (Hordeum vulgare) east of the Fertile Crescent. Proc. Natl. Acad. Sci. USA 104, 3289–3294 (2007).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  27. van Zeist, W. & Bakker-Heeres, J.A.H. Archaeological studies in the Levant. 1. Neolithic sites in the Damascus basin: Aswad, Ghoraifé, Ramad. Palaeohistoria 24, 165–256 (1985).

    Google Scholar 

  28. Snir, A. et al. The origin of cultivation and proto-weeds, long before Neolithic farming. PLoS One 10, e0131422 (2015).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  29. Abdel-Ghani, A.H., Parzies, H.K., Omary, A. & Geiger, H.H. Estimating the outcrossing rate of barley landraces and wild barley populations collected from ecologically different regions of Jordan. Theor. Appl. Genet. 109, 588–595 (2004).

    Article  PubMed  Google Scholar 

  30. Russell, J. et al. 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 Phytol. 191, 564–578 (2011).

    Article  PubMed  Google Scholar 

  31. Alexander, D.H., Novembre, J. & Lange, K. Fast model-based estimation of ancestry in unrelated individuals. Genome Res. 19, 1655–1664 (2009).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  32. Durand, E.Y., Patterson, N., Reich, D. & Slatkin, M. Testing for ancient admixture between closely related populations. Mol. Biol. Evol. 28, 2239–2252 (2011).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  33. Jakob, S.S. & Blattner, F.R. A chloroplast genealogy of Hordeum (Poaceae): long-term persisting haplotypes, incomplete lineage sorting, regional extinction, and the consequences for phylogenetic inference. Mol. Biol. Evol. 23, 1602–1612 (2006).

    Article  CAS  PubMed  Google Scholar 

  34. Dickson, A.D. et al. Barley: Origin, Botany, Culture, Winter Hardiness, Genetics, Utilization, Pests (US Science and Education Administration, US Department of Agriculture, 1978).

  35. Russell, J. et al. Genetic diversity and ecological niche modelling of wild barley: refugia, large-scale post-LGM range expansion and limited mid-future climate threats? PLoS One 9, e86021 (2014).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  36. Frumin, S., Maeir, A.M., Kolska Horwitz, L. & Weiss, E. Studying ancient anthropogenic impacts on current floral biodiversity in the Southern Levant as reflected by the Philistine migration. Sci. Rep. 5, 13308 (2015).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  37. Meiri, M. et al. Ancient DNA and population turnover in Southern Levantine pigs—signature of the Sea Peoples migration? Sci. Rep. 3, 3035 (2013).

    Article  PubMed  PubMed Central  Google Scholar 

  38. Giuffra, E. et al. The origin of the domestic pig: independent domestication and subsequent introgression. Genetics 154, 1785–1791 (2000).

    CAS  PubMed  PubMed Central  Google Scholar 

  39. Haber, M. et al. Genome-wide diversity in the Levant reveals recent structuring by culture. PLoS Genet. 9, e1003316 (2013).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  40. Bilgic, H., Hakki, E.E., Pandey, A., Khan, M.K. & Akkaya, M.S. Ancient DNA from 8400 year-old Çatalhöyük wheat: implications for the origin of Neolithic agriculture. PLoS One 11, e0151974 (2016).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  41. Arend, D. et al. e!DAL—a framework to store, share and publish research data. BMC Bioinformatics 15, 214 (2014).

    Article  PubMed  PubMed Central  Google Scholar 

  42. Schick, T. The early basketry and textiles from caves in the northern Judean desert. Part II'Atiqot 41, 223–239 (2002).

    Google Scholar 

  43. Kistler, L. Ancient DNA extraction from plants. Methods Mol. Biol. 840, 71–79 (2012).

    Article  CAS  PubMed  Google Scholar 

  44. Meyer, M. & Kircher, M. Illumina sequencing library preparation for highly multiplexed target capture and sequencing. Cold Spring Harb. Protoc. 2010, prot5448 (2010).

    Article  Google Scholar 

  45. Kircher, M., Sawyer, S. & Meyer, M. Double indexing overcomes inaccuracies in multiplex sequencing on the Illumina platform. Nucleic Acids Res. 40, e3 (2012).

    Article  CAS  PubMed  Google Scholar 

  46. Himmelbach, A., Knauft, M. & Stein, N. Plant sequence capture optimised for Illumina sequencing. Bio Protoc. 4, e1166 (2014).

    Article  Google Scholar 

  47. Kircher, M. Analysis of high-throughput ancient DNA sequencing data. Methods Mol. Biol. 840, 197–228 (2012).

    Article  CAS  PubMed  Google Scholar 

  48. Renaud, G., Stenzel, U. & Kelso, J. leeHom: adaptor trimming and merging for Illumina sequencing reads. Nucleic Acids Res. 42, e141 (2014).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  49. Gallego Llorente, M. et al. Ancient Ethiopian genome reveals extensive Eurasian admixture throughout the African continent. Science 350, 820–822 (2015).

    Article  CAS  PubMed  Google Scholar 

  50. International Barley Genome Sequencing Consortium. A physical, genetic and functional sequence assembly of the barley genome. Nature 491, 711–716 (2012).

  51. Saski, C. et al. Complete chloroplast genome sequences of Hordeum vulgare, Sorghum bicolor and Agrostis stolonifera, and comparative analyses with other grass genomes. Theor. Appl. Genet. 115, 571–590 (2007).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  52. Li, H. Aligning sequence reads, clone sequences and assembly contigs with BWA-MEM. Preprint at https://arxiv.org/abs/1303.3997 (2013).

  53. Li, H. et al. The Sequence Alignment/Map format and SAMtools. Bioinformatics 25, 2078–2079 (2009).

    PubMed  PubMed Central  Google Scholar 

  54. Jónsson, H., Ginolhac, A., Schubert, M., Johnson, P.L. & Orlando, L. mapDamage2.0: fast approximate Bayesian estimates of ancient DNA damage parameters. Bioinformatics 29, 1682–1684 (2013).

    Article  PubMed  PubMed Central  CAS  Google Scholar 

  55. Li, H. A statistical framework for SNP calling, mutation discovery, association mapping and population genetical parameter estimation from sequencing data. Bioinformatics 27, 2987–2993 (2011).

    CAS  PubMed  PubMed Central  Google Scholar 

  56. Quinlan, A.R. & Hall, I.M. BEDTools: a flexible suite of utilities for comparing genomic features. Bioinformatics 26, 841–842 (2010).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  57. Zheng, X. et al. A high-performance computing toolset for relatedness and principal component analysis of SNP data. Bioinformatics 28, 3326–3328 (2012).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  58. Jakobsson, M. & Rosenberg, N.A. CLUMPP: a cluster matching and permutation program for dealing with label switching and multimodality in analysis of population structure. Bioinformatics 23, 1801–1806 (2007).

    Article  CAS  PubMed  Google Scholar 

  59. Patterson, N. et al. Ancient admixture in human history. Genetics 192, 1065–1093 (2012).

    Article  PubMed  PubMed Central  Google Scholar 

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Acknowledgements

We gratefully acknowledge B. Schäfer for providing photographs of barley spikes and A. Fiebig for sequence data submission. This work was supported by a grant of the Israel Science Foundation (1179/13) to E.W., funding from the Endowed Chair in Molecular Genetics Applied to Crop Improvement at the University of Minnesota and the Triticeae Coordinated Agricultural Project, USDA-NIFA 2011-68002-30029 to G.J.M., and core funding of IPK Gatersleben to N.S. and M.M. R.W., J.R. and I.K.D. were supported by Research Programme funding from the Scottish government and the University of Dundee (R.W.). N.M. and U.D. acknowledge a grant from the Irene Levi Sela CARE Foundation.

Author information

Author notes

  1. Martin Mascher, Verena J Schuenemann & Benjamin Kilian

    Present address: Present address: Bayer CropScience, BCS Breeding and Trait Development, Zwijnaarde (Gent), Belgium.,

  2. These authors contributed equally to this work.

Authors and Affiliations

  1. Leibniz Institute of Plant Genetics and Crop Plant Research (IPK) Gatersleben, Seeland, Germany

    Martin Mascher, Axel Himmelbach, Mona Schreiber, Benjamin Kilian & Nils Stein

  2. German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Leipzig, Germany

    Martin Mascher

  3. Institute for Archaeological Sciences, University of Tübingen, Tübingen, Germany

    Verena J Schuenemann, Ella Reiter, Simone Riehl & Johannes Krause

  4. Senckenberg Center for Human Evolution and Paleoenvironment, University of Tübingen, Tübingen, Germany

    Verena J Schuenemann, Simone Riehl & Johannes Krause

  5. Institute of Archaeology, Hebrew University, Jerusalem, Israel

    Uri Davidovich

  6. Laboratory of Archaeozoology, Zinman Institute of Archaeology, University of Haifa, Haifa, Israel

    Nimrod Marom

  7. Department of Botany, University of British Columbia, Vancouver, British Columbia, Canada

    Sariel Hübner

  8. Department of Biotechnology, Tel Hai College, Upper Galilee, Israel

    Sariel Hübner

  9. Institute of Evolution, University of Haifa, Haifa, Israel

    Abraham Korol & Tzion Fahima

  10. Department of Evolutionary and Environmental Biology, University of Haifa, Haifa, Israel

    Abraham Korol & Tzion Fahima

  11. Martin (Szusz) Department of Land of Israel Studies and Archaeology, Bar-Ilan University, Ramat-Gan, Israel

    Michal David & Ehud Weiss

  12. Department of Biomolecular Engineering, University of California, Santa Cruz, Santa Cruz, California, USA.,

    Samuel H Vohr & Richard E Green

  13. Cell and Molecular Sciences, James Hutton Institute, Invergowrie, Dundee, UK

    Ian K Dawson, Joanne Russell & Robbie Waugh

  14. Department of Plant Biology, University of Minnesota, St. Paul, Minnesota, USA

    Gary J Muehlbauer

  15. Department of Agronomy and Plant Genetics, University of Minnesota, St. Paul, Minnesota, USA

    Gary J Muehlbauer

  16. Division of Plant Sciences, University of Dundee, Dundee, UK

    Robbie Waugh

  17. Max Planck Institute for the Science of Human History, Jena, Germany

    Johannes Krause

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  1. Martin Mascher
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Contributions

E.W., T.F., N.S. and J.K. conceived the study. E.W., T.F., N.S., J.K., V.J.S. and M.M. designed experiments. N.M., U.D., M.D., S.R. and E.W. performed excavations and archaeobotanical analyses. V.J.S., A.H. and E.R. performed the ancient DNA experiments. M.M., S.H., A.K., M.S., S.H.V. and R.E.G. analyzed data. J.R., M.M., I.K.D., B.K., G.J.M., N.S. and R.W. provided exome capture data. M.M., V.J.S., A.H., S.R., T.F., J.K., E.W. and N.S. wrote the manuscript with input from all co-authors. All authors read and approved the manuscript.

Corresponding authors

Correspondence to Tzion Fahima, Johannes Krause, Ehud Weiss or Nils Stein.

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The authors declare no competing financial interests.

Integrated supplementary information

Supplementary Figure 1 Plan of Yoram Cave—top plan and sections.

The arrow indicates the entrance to the cave and the human-made wall across it. Note the boulders in the northern room and the relatively horizontal surface in the southern room. Gray shaded borders indicate that the edge of the cave is cut into rock.

Supplementary Figure 2 Photographs from the excavation.

(a) Masada Southern Cave complex, with three caves located in the southeastern cliff of the Masada Horst. The dotted red line marks the easiest trail giving access to the caves. (b) The entrance of Yoram Cave, facing southeast, in an almost vertical cliff, some 4 m above the trail leading to the cave. (c) The south room during excavation of locus 3.

Supplementary Figure 3 Photographs of ancient barley grains used for DNA extraction.

Supplementary Figure 4 Length distribution of sequence fragments.

Supplementary Figure 5 Nucleotide misincorporation profiles in reads mapped to the whole-genome shotgun assembly of barley cv. Morex.

The proportion of C>T misincorporation sites (red) is compared to the G>A baseline (black).

Supplementary Figure 6 Nucleotide misincorporation profiles in reads mapped to the barley chloroplast genome assembly.

The proportion of C>T substitutions (red) is compared to the G>A baseline (black).

Supplementary Figure 7 Haplotype of Btr1, Btr2 and Vrs1.

The contigs of the Morex whole-genome sequencing assembly harboring these genes were identified by BLAST searches. Read depth in the deeply sequenced sample JK3014 is shown in black. Position on the respective whole-genome sequencing contig is indicated along the upper axis. Distance (in genomic sequence) from the start codon of each gene is shown along the bottom axis. Btr1 and Btr2 are single-exon genes. Gray bars indicate the positions of the three exons of Vrs1. The positions of SNPs are highlighted by vertical lines. The Morex allele is shown above and the JK3014 allele is shown below the lines. SNPs with a previously reported functional effect are shown in red. JK3014 carries a loss-of-function allele of Btr1, whereas Btr2 and Vrs1 have wild-type alleles. The coding sequences of Btr1 and Btr2 are identical to those in the haplotype of cv. Haruna Nijo as reported by Komatsuda et al. (NCBI GenBank accession KR813337.1). The sequence of Vrs1 matches the Vrs1.b2 allele as designated by Komatsuda et al.22.

Supplementary Figure 8 Relationship between genetic similarity and geographical distance.

(a–f) Scatterplots of genetic similarity and geographical distance between 91 extant wild barley accessions sampled across the Fertile Crescent and archaeological sample JK3014 found at Yoram Cave and sequenced to higher depth using all SNPs (a), a two-rowed cultivated landrace from Israel (b), a two-rowed cultivated landrace from Egypt (c), the ancient sample JK3014 found at Yoram Cave and sequenced to higher depth excluding transition SNPs (d), two-rowed cultivated landraces from Israel excluding transition SNPs (e) and two-rowed cultivated landraces from Egypt excluding transition SNPs (f). The geographical position attributed to each sample is as follows: 31.3141° N, 35.353° E for a and d, 31.7156° N, 35.1871° E for b and e, and 31.193° N, 29.904° E for c and f. Correlation coefficients and P values for the geographically proximate and distant subsets are indicated in blue and red, respectively.

Supplementary Figure 9 Cross-validation error of ADMIXTURE analysis.

Box plots were calculated from 20 replicate runs for each K value.

Supplementary Figure 10 ADMIXTURE analysis with K = 5 for domesticated and wild samples.

Top, domesticated samples; bottom, wild samples. Colors in both panels correspond to the same ancestral populations. Sample names and countries of origin are indicated above and below the plots, respectively. Ancient samples are highlighted by blue borders.

Supplementary information

Supplementary Text and Figures

Supplementary Figures 1–10 and Supplementary Tables 1, 2 and 4. (PDF 1168 kb)

Supplementary Table 3

Identity-by-state matrix. (XLSX 26 kb)

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Mascher, M., Schuenemann, V., Davidovich, U. et al. Genomic analysis of 6,000-year-old cultivated grain illuminates the domestication history of barley. Nat Genet 48, 1089–1093 (2016). https://doi.org/10.1038/ng.3611

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