Structural variation in the pangenome of wild and domesticated barley
2 Carlsberg Research Laboratory
3 IPSIM - Institut des Sciences des Plantes de Montpellier
4 The James Hutton Institute
5 UZH - Universität Zürich [Zürich] = University of Zurich
6 Agriculture and Agri-Food Canada, Saskatoon Research Centre
7 Agriculture and Agri-Food Canada
8 UBC - University of British Columbia [Canada]
9 DSMZ - Leibniz-Institut DSMZ-Deutsche Sammlung von Mikroorganismen und Zellkulturen GmbH / Leibniz Institute DSMZ-German Collection of Microorganisms and Cell Cultures
10 School of Agriculture, Food and Wine
11 Murdoch University [Perth]
12 HMGU - Helmholtz Zentrum München = German Research Center for Environmental Health
13 Lund University
14 GDAAS - Guangdong Academy of Agricultural Sciences
15 KDRI - Kazusa DNA Research Institute
16 La Trobe University
17 KAUST - King Abdullah University of Science and Technology [Saudi Arabia]
18 University of Adelaide
19 Government of Western Australia
20 Yangtze University
21 Martin-Luther-University Halle-Wittenberg
22 TUM - Technische Universität Munchen - Technical University Munich - Université Technique de Munich
23 UMN - University of Minnesota [Twin Cities]
24 JKI - Julius Kühn-Institut - Federal Research Centre for Cultivated Plants
25 Secobra Recherches
26 U of S - University of Saskatchewan [Saskatoon, Canada]
27 Tottori University
28 Okayama University
29 University of Dundee
30 iDiv - German Centre for Integrative Biodiversity Research
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Résumé
P an genomes are collections of annotated genome sequences of multiple individuals of a species 1 . The structural variants uncovered by these datasets are a major asset to genetic analysis in crop plants 2 . Here we report a pangenome of barley comprising long-read sequence assemblies of 76 wild and domesticated genomes and short-read sequence data of 1,315 genotypes. An expanded catalogue of sequence variation in the crop includes structurally complex loci that are rich in gene copy number variation. To demonstrate the utility of the pangenome, we focus on four loci involved in disease resistance, plant architecture, nutrient release and trichome development. Novel allelic variation at a powdery mildew resistance locus and population-specific copy number gains in a regulator of vegetative branching were found. Expansion of a family of starch-cleaving enzymes in elite malting barleys was linked to shifts in enzymatic activity in micro-malting trials. Deletion of an enhancer motif is likely to change the developmental trajectory of the hairy appendages on barley grains. Our findings indicate that allelic diversity at structurally complex loci may have helped crop plants to adapt to new selective regimes in agricultural ecosystems.
Reliable crop yields fuelled the rise of human civilizations. As people embraced a new way of life, cultivated plants, too, had to adapt to the needs of their domesticators. There are different adaptive requirements in a wild compared with an arable habitat. Crop plants and their wild progenitors differ in how many vegetative branches they initiate or how many seeds or fruits they produce and when. A case in point is barley (Hordeum vulgare): in six-rowed forms of the crops, thrice as many grains set as in the ancestral two-rowed forms. This change was brought about by knockout mutations 3 of a recently evolved regulator 4 of inflorescence development. Consequently, six-rowed barleys came to predominate in most barley-growing regions 5 . Taking a broader view of the environment as a set of exogeneous factors that drive natural selection, barley provides another fascinating, and economically important, example. The process of malting involves the sprouting of moist barley grains, driving the release of enzymes that break down starch into fermentable sugars. In the wild, various environmental cues can trigger germination to improve the odds of the emerging seedling encountering favourable weather conditions for subsequent growth 6 . In the malt house, by contrast, germination has to be fast and uniform in modern cultivars to satisfy the desired specifications of the industry. In addition to these examples, traits such as disease resistance, plant architecture and nutrient use have been a focus for plant breeders and studied intensively by barley geneticists 7 . Although barley genetic analysis flourished during a 'classical' period 8 in the first half of the 20th century, it started to lag behind small-genome models because of difficulties in adapting molecular biology techniques to a large genome rich in repeats 9 . However, interest in barley as a diploid model for temperate cereals has surged again as DNA sequencing became more powerful. High-quality sequences of several barley genomes have been recently assembled 10 . New sequencing technologies have shifted the focus of
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