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      Analysis of the bread wheat genome using whole genome shotgun sequencing

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          Summary

          Bread wheat ( Triticum aestivum) is a globally important crop, accounting for 20% of the calories consumed by mankind. We sequenced its large and challenging 17 Gb hexaploid genome using 454 pyrosequencing and compared this with the sequences of diploid ancestral and progenitor genomes. Between 94,000-96,000 genes were identified, and two-thirds were assigned to the A, B and D genomes. High-resolution synteny maps identified many small disruptions to conserved gene order. We show the hexaploid genome is highly dynamic, with significant loss of gene family members upon polyploidization and domestication, and an abundance of gene fragments. Several classes of genes involved in energy harvesting, metabolism and growth are among expanded gene families that could be associated with crop productivity. Our analyses, coupled with the identification of extensive genetic variation, provide a new resource for accelerating gene discovery and improving this major crop.

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          Most cited references29

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          The Sorghum bicolor genome and the diversification of grasses.

          Sorghum, an African grass related to sugar cane and maize, is grown for food, feed, fibre and fuel. We present an initial analysis of the approximately 730-megabase Sorghum bicolor (L.) Moench genome, placing approximately 98% of genes in their chromosomal context using whole-genome shotgun sequence validated by genetic, physical and syntenic information. Genetic recombination is largely confined to about one-third of the sorghum genome with gene order and density similar to those of rice. Retrotransposon accumulation in recombinationally recalcitrant heterochromatin explains the approximately 75% larger genome size of sorghum compared with rice. Although gene and repetitive DNA distributions have been preserved since palaeopolyploidization approximately 70 million years ago, most duplicated gene sets lost one member before the sorghum-rice divergence. Concerted evolution makes one duplicated chromosomal segment appear to be only a few million years old. About 24% of genes are grass-specific and 7% are sorghum-specific. Recent gene and microRNA duplications may contribute to sorghum's drought tolerance.
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            Differentiation of the maize subgenomes by genome dominance and both ancient and ongoing gene loss.

            Ancient tetraploidies are found throughout the eukaryotes. After duplication, one copy of each duplicate gene pair tends to be lost (fractionate). For all studied tetraploidies, the loss of duplicated genes, known as homeologs, homoeologs, ohnologs, or syntenic paralogs, is uneven between duplicate regions. In maize, a species that experienced a tetraploidy 5-12 million years ago, we show that in addition to uneven ancient gene loss, the two complete genomes contained within maize are differentiated by ongoing fractionation among diverse inbreds as well as by a pattern of overexpression of genes from the genome that has experienced less gene loss. These expression differences are consistent over a range of experiments quantifying RNA abundance in different tissues. We propose that the universal bias in gene loss between the genomes of this ancient tetraploid, and perhaps all tetraploids, is the result of selection against loss of the gene responsible for the majority of total expression for a duplicate gene pair. Although the tetraploidy of maize is ancient, biased gene loss and expression continue today and explain, at least in part, the remarkable genetic diversity found among modern maize cultivars.
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              Pack-MULE transposable elements mediate gene evolution in plants.

              Mutator-like transposable elements (MULEs) are found in many eukaryotic genomes and are especially prevalent in higher plants. In maize, rice and Arabidopsis a few MULEs were shown to carry fragments of cellular genes. These chimaeric elements are called Pack-MULEs in this study. The abundance of MULEs in rice and the availability of most of the genome sequence permitted a systematic analysis of the prevalence and nature of Pack-MULEs in an entire genome. Here we report that there are over 3,000 Pack-MULEs in rice containing fragments derived from more than 1,000 cellular genes. Pack-MULEs frequently contain fragments from multiple chromosomal loci that are fused to form new open reading frames, some of which are expressed as chimaeric transcripts. About 5% of the Pack-MULEs are represented in collections of complementary DNA. Functional analysis of amino acid sequences and proteomic data indicate that some captured gene fragments might be functional. Comparison of the cellular genes and Pack-MULE counterparts indicates that fragments of genomic DNA have been captured, rearranged and amplified over millions of years. Given the abundance of Pack-MULEs in rice and the widespread occurrence of MULEs in all characterized plant genomes, gene fragment acquisition by Pack-MULEs might represent an important new mechanism for the evolution of genes in higher plants.
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                Author and article information

                Journal
                0410462
                6011
                Nature
                Nature
                Nature
                0028-0836
                1476-4687
                15 October 2012
                29 November 2012
                29 May 2013
                : 491
                : 7426
                : 705-710
                Affiliations
                [1 ]Centre for Genome Research, University of Liverpool, Liverpool, UK
                [2 ]MIPS/IBIS, Helmholtz-Zentrum München, Neuherberg, DE
                [3 ]School of Biological Sciences, University of Bristol, Bristol, UK
                [4 ]John Innes Centre, Norwich, UK
                [5 ]Cold Spring Harbor Laboratory, Cold Spring Harbor, NY, USA
                [6 ]European Bioinformatics Institute, Hinxton, UK
                [7 ]USDA Western Regional Laboratory, Albany, CA, USA
                [8 ]Dept Agronomy and Range Science, UC Davis, Davis, CA, USA
                [9 ]Dept Plant Pathology, Kansas State University, Manhattan, KS, USA.
                Author notes
                [§]

                joint first authors

                [* ]senior and corresponding authors

                Requests for materials should be addressed to Michael Bevan, and correspondence to Klaus Mayer, Michael Bevan, Neil Hall or Keith Edwards.

                Author Contributions K.J.E., M.W.B., N.H. and A.H. conceived the project, R.M., M.K., M.T., I.B., J.D., M.C.L., O.A., S.K., N.Huo, B.G., S.S., provided data and advice, R.dA., N.McK. and S.K. conducted experiments, K.F.X.M., N.H. and M.W.B. planned and conducted analyses, R.B., M.S., M.P., G.B., A.A., D.B., D.W., P.K., A.H. carried out analyses. K.J.E., A.H., R.M., R.B., contributed to the text and M.W.B., N.H., and K.F.X.M. wrote the manuscript. All authors commented on the manuscript.

                Article
                EMS50090
                10.1038/nature11650
                3510651
                23192148
                36a98bea-f00e-408d-992a-09b6d5f5be44

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                wheat/polyploid/genome,analysis/food,security/next-generation,sequencing
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                wheat/polyploid/genome, analysis/food, security/next-generation, sequencing

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