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      The genome of cowpea ( Vigna unguiculata [L.] Walp.)

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      1 , , 2 , 14 , , 1 , 3 , 2 , 2 , 4 , 5 , 6 , 4 , 5 , 6 , 7 , 7 , 1 , 1 , 1 , 8 , 9 , 9 , 10 , 9 , 11 , 11 , 12 , 12 , 13 , 2
      The Plant Journal
      John Wiley and Sons Inc.
      chromosomal inversion, cowpea, domestication, genome annotation, genome evolution, genome size, next‐generation sequencing, legumes, Phaseolus vulgaris, repetitive elements, Vigna unguiculata

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          Summary

          Cowpea ( Vigna unguiculata [L.] Walp.) is a major crop for worldwide food and nutritional security, especially in sub‐Saharan Africa, that is resilient to hot and drought‐prone environments. An assembly of the single‐haplotype inbred genome of cowpea IT97K‐499‐35 was developed by exploiting the synergies between single‐molecule real‐time sequencing, optical and genetic mapping, and an assembly reconciliation algorithm. A total of 519 Mb is included in the assembled sequences. Nearly half of the assembled sequence is composed of repetitive elements, which are enriched within recombination‐poor pericentromeric regions. A comparative analysis of these elements suggests that genome size differences between Vigna species are mainly attributable to changes in the amount of Gypsy retrotransposons. Conversely, genes are more abundant in more distal, high‐recombination regions of the chromosomes; there appears to be more duplication of genes within the NBS‐LRR and the SAUR‐like auxin superfamilies compared with other warm‐season legumes that have been sequenced. A surprising outcome is the identification of an inversion of 4.2 Mb among landraces and cultivars, which includes a gene that has been associated in other plants with interactions with the parasitic weed Striga gesnerioides. The genome sequence facilitated the identification of a putative syntelog for multiple organ gigantism in legumes. A revised numbering system has been adopted for cowpea chromosomes based on synteny with common bean ( Phaseolus vulgaris). An estimate of nuclear genome size of 640.6 Mbp based on cytometry is presented.

          Significance Statement

          State‐of‐the‐art technologies and assembly methods were used to generate a reference genome sequence of cowpea, a drought‐resilient crop on which millions of people in sub‐Saharan Africa depend as a source of protein. This sequence facilitated the identification of: repetitive elements and gene families expanded in cowpea compared with other closely related legumes; a large and apparently rare chromosomal inversion; and an interesting candidate gene that is associated with several domestication‐related traits.

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          The roles of segmental and tandem gene duplication in the evolution of large gene families in Arabidopsis thaliana

          Background Most genes in Arabidopsis thaliana are members of gene families. How do the members of gene families arise, and how are gene family copy numbers maintained? Some gene families may evolve primarily through tandem duplication and high rates of birth and death in clusters, and others through infrequent polyploidy or large-scale segmental duplications and subsequent losses. Results Our approach to understanding the mechanisms of gene family evolution was to construct phylogenies for 50 large gene families in Arabidopsis thaliana, identify large internal segmental duplications in Arabidopsis, map gene duplications onto the segmental duplications, and use this information to identify which nodes in each phylogeny arose due to segmental or tandem duplication. Examples of six gene families exemplifying characteristic modes are described. Distributions of gene family sizes and patterns of duplication by genomic distance are also described in order to characterize patterns of local duplication and copy number for large gene families. Both gene family size and duplication by distance closely follow power-law distributions. Conclusions Combining information about genomic segmental duplications, gene family phylogenies, and gene positions provides a method to evaluate contributions of tandem duplication and segmental genome duplication in the generation and maintenance of gene families. These differences appear to correspond meaningfully to differences in functional roles of the members of the gene families.
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            LTRharvest, an efficient and flexible software for de novo detection of LTR retrotransposons

            Background Transposable elements are abundant in eukaryotic genomes and it is believed that they have a significant impact on the evolution of gene and chromosome structure. While there are several completed eukaryotic genome projects, there are only few high quality genome wide annotations of transposable elements. Therefore, there is a considerable demand for computational identification of transposable elements. LTR retrotransposons, an important subclass of transposable elements, are well suited for computational identification, as they contain long terminal repeats (LTRs). Results We have developed a software tool LTRharvest for the de novo detection of full length LTR retrotransposons in large sequence sets. LTRharvest efficiently delivers high quality annotations based on known LTR transposon features like length, distance, and sequence motifs. A quality validation of LTRharvest against a gold standard annotation for Saccharomyces cerevisae and Drosophila melanogaster shows a sensitivity of up to 90% and 97% and specificity of 100% and 72%, respectively. This is comparable or slightly better than annotations for previous software tools. The main advantage of LTRharvest over previous tools is (a) its ability to efficiently handle large datasets from finished or unfinished genome projects, (b) its flexibility in incorporating known sequence features into the prediction, and (c) its availability as an open source software. Conclusion LTRharvest is an efficient software tool delivering high quality annotation of LTR retrotransposons. It can, for example, process the largest human chromosome in approx. 8 minutes on a Linux PC with 4 GB of memory. Its flexibility and small space and run-time requirements makes LTRharvest a very competitive candidate for future LTR retrotransposon annotation projects. Moreover, the structured design and implementation and the availability as open source provides an excellent base for incorporating novel concepts to further improve prediction of LTR retrotransposons.
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              Arabidopsis cytokinin receptor mutants reveal functions in shoot growth, leaf senescence, seed size, germination, root development, and cytokinin metabolism.

              We used loss-of-function mutants to study three Arabidopsis thaliana sensor histidine kinases, AHK2, AHK3, and CRE1/AHK4, known to be cytokinin receptors. Mutant seeds had more rapid germination, reduced requirement for light, and decreased far-red light sensitivity, unraveling cytokinin functions in seed germination control. Triple mutant seeds were more than twice as large as wild-type seeds. Genetic analysis indicated a cytokinin-dependent endospermal and/or maternal control of embryo size. Unchanged red light sensitivity of mutant hypocotyl elongation suggests that previously reported modulation of red light signaling by A-type response regulators may not depend on cytokinin. Combined loss of AHK2 and AHK3 led to the most prominent changes during vegetative development. Leaves of ahk2 ahk3 mutants formed fewer cells, had reduced chlorophyll content, and lacked the cytokinin-dependent inhibition of dark-induced chlorophyll loss, indicating a prominent role of AHK2 and, particularly, AHK3 in the control of leaf development. ahk2 ahk3 double mutants developed a strongly enhanced root system through faster growth of the primary root and, more importantly, increased branching. This result supports a negative regulatory role for cytokinin in root growth regulation. Increased cytokinin content of receptor mutants indicates a homeostatic control of steady state cytokinin levels through signaling. Together, the analyses reveal partially redundant functions of the cytokinin receptors and prominent roles for the AHK2/AHK3 receptor combination in quantitative control of organ growth in plants, with opposite regulatory functions in roots and shoots.
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                Author and article information

                Contributors
                stelo@cs.ucr.edu
                maria.munoz_amatriain@colostate.edu
                Journal
                Plant J
                Plant J
                10.1111/(ISSN)1365-313X
                TPJ
                The Plant Journal
                John Wiley and Sons Inc. (Hoboken )
                0960-7412
                1365-313X
                28 May 2019
                June 2019
                : 98
                : 5 ( doiID: 10.1111/tpj.2019.98.issue-5 )
                : 767-782
                Affiliations
                [ 1 ] Department of Computer Science and Engineering University of California Riverside CA 92521 USA
                [ 2 ] Department of Botany and Plant Sciences University of California Riverside CA 92521 USA
                [ 3 ] US Department of Energy Joint Genome Institute Walnut Creek CA 94598 USA
                [ 4 ] Natural Resources Institute Finland (Luke) Helsinki Finland
                [ 5 ] Institute of Biotechnology University of Helsinki Helsinki Finland
                [ 6 ] Viikki Plant Science Centre University of Helsinki Helsinki Finland
                [ 7 ] Department of Plant Sciences University of California Davis CA 95616 USA
                [ 8 ] Institut de Recherche en Horticulture et Semences INRA Université d'Angers 49071 Beaucouzé France
                [ 9 ] Department of Nematology University of California Riverside CA 92521 USA
                [ 10 ] Departamento de Fitopatologia Instituto de Ciências Biológicas Universidade de Brasília Brasília DF Brazil
                [ 11 ] Centre of the Region Haná for Biotechnological and Agricultural Research Institute of Experimental Botany Olomouc Czech Republic
                [ 12 ] National Center for Genome Resources Santa Fe NM 87505 USA
                [ 13 ] US Department of Agriculture–Agricultural Research Service Ames IA USA
                [ 14 ]Present address: Department of Soil and Crop Sciences Colorado State University Fort Collins CO 80523 USA
                Author notes
                [*] [* ]For correspondence (e‐mails stelo@ 123456cs.ucr.edu and maria.munoz_amatriain@ 123456colostate.edu ).
                [†]

                These authors contributed equally to this work.

                Author information
                https://orcid.org/0000-0002-2696-7274
                https://orcid.org/0000-0002-4476-1691
                https://orcid.org/0000-0002-6263-0492
                https://orcid.org/0000-0002-9759-3775
                Article
                TPJ14349
                10.1111/tpj.14349
                6852540
                31017340
                802be6ae-397a-4967-a495-14e6bdbfbabf
                © 2019 The Authors. The Plant Journal published by John Wiley & Sons Ltd and Society for Experimental Biology.

                This is an open access article under the terms of the http://creativecommons.org/licenses/by/4.0/ License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

                History
                : 28 January 2019
                : 28 March 2019
                : 25 March 2019
                Page count
                Figures: 3, Tables: 1, Pages: 16, Words: 13349
                Funding
                Funded by: NSF , open-funder-registry 10.13039/100000001;
                Award ID: IOS‐1543963
                Award ID: IIS‐1526742
                Award ID: IIS‐1814359
                Funded by: Czech Ministry of Education, Youth and Sports
                Award ID: LO1204
                Funded by: Agricultural Research Service , open-funder-registry 10.13039/100007917;
                Award ID: 5030‐21000‐069‐00‐D
                Funded by: Academy of Finland “Papugeno” , open-funder-registry 10.13039/501100002341;
                Award ID: 298314
                Categories
                Resource
                Resource
                Custom metadata
                2.0
                June 2019
                Converter:WILEY_ML3GV2_TO_JATSPMC version:5.7.1 mode:remove_FC converted:13.11.2019

                Plant science & Botany
                chromosomal inversion,cowpea,domestication,genome annotation,genome evolution,genome size,next‐generation sequencing,legumes,phaseolus vulgaris,repetitive elements,vigna unguiculata

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