title: The genome of the seagrass Zostera marina reveals angiosperm adaptation to the sea.
abstract: Seagrasses colonized the sea on at least three independent occasions to form the basis of one of the most productive and widespread coastal ecosystems on the planet. Here we report the genome of Zostera marina (L.), the first, to our knowledge, marine angiosperm to be fully sequenced. This reveals unique insights into the genomic losses and gains involved in achieving the structural and physiological adaptations required for its marine lifestyle, arguably the most severe habitat shift ever accomplished by flowering plants. Key angiosperm innovations that were lost include the entire repertoire of stomatal genes, genes involved in the synthesis of terpenoids and ethylene signalling, and genes for ultraviolet protection and phytochromes for far-red sensing. Seagrasses have also regained functions enabling them to adjust to full salinity. Their cell walls contain all of the polysaccharides typical of land plants, but also contain polyanionic, low-methylated pectins and sulfated galactans, a feature shared with the cell walls of all macroalgae and that is important for ion homoeostasis, nutrient uptake and O2/CO2 exchange through leaf epidermal cells. The Z. marina genome resource will markedly advance a wide range of functional ecological studies from adaptation of marine ecosystems under climate warming, to unravelling the mechanisms of osmoregulation under high salinities that may further inform our understanding of the evolution of salt tolerance in crop plants.
authors.affiliation: Groningen Institute of Evolutionary Life Sciences (GELIFES), University of Groningen, PO Box 11103, 9700 CC Groningen, The Netherlands.
authors.forename: Jeanine L
authors.initials: JL
authors.lastname: Olsen
authors.affiliation: Department of Plant Systems Biology, VIB and Department of Plant Biotechnology and Bioinformatics, Ghent University, Technologiepark 927, B-9052 Ghent, Belgium.
authors.forename: Pierre
authors.initials: P
authors.lastname: Rouzé
authors.affiliation: Department of Plant Systems Biology, VIB and Department of Plant Biotechnology and Bioinformatics, Ghent University, Technologiepark 927, B-9052 Ghent, Belgium.
authors.forename: Bram
authors.initials: B
authors.lastname: Verhelst
authors.affiliation: Department of Plant Systems Biology, VIB and Department of Plant Biotechnology and Bioinformatics, Ghent University, Technologiepark 927, B-9052 Ghent, Belgium.
authors.forename: Yao-Cheng
authors.initials: YC
authors.lastname: Lin
authors.affiliation: GEOMAR Helmholtz Centre for Ocean Research-Kiel, Evolutionary Ecology, Düsternbrooker Weg 20, D-24105 Kiel, Germany.
authors.forename: Till
authors.initials: T
authors.lastname: Bayer
authors.affiliation: Sorbonne Université, UPMC Univ Paris 06, CNRS, UMR 8227, Integrative Biology of Marine Models, Station Biologique de Roscoff, CS 90074, F-29688, Roscoff cedex, France.
authors.forename: Jonas
authors.initials: J
authors.lastname: Collen
authors.affiliation: Stazione Zoologica Anton Dohrn, Villa Comunale, 80121 Naples, Italy.
authors.forename: Emanuela
authors.initials: E
authors.lastname: Dattolo
authors.affiliation: Dipartimento di Scienze Agrarie e Ambientali, University of Udine, Via delle Scienze 206, 33100 Udine, Italy.
authors.forename: Emanuele
authors.initials: E
authors.lastname: De Paoli
authors.affiliation: Sorbonne Université, UPMC Univ Paris 06, CNRS, UMR 8227, Integrative Biology of Marine Models, Station Biologique de Roscoff, CS 90074, F-29688, Roscoff cedex, France.
authors.forename: Simon
authors.initials: S
authors.lastname: Dittami
authors.affiliation: INRA, UR1164 URGI-Research Unit in Genomics-Info, INRA de Versailles-Grignon, Route de Saint-Cyr, Versailles 78026, France.
authors.forename: Florian
authors.initials: F
authors.lastname: Maumus
authors.affiliation: Sorbonne Université, UPMC Univ Paris 06, CNRS, UMR 8227, Integrative Biology of Marine Models, Station Biologique de Roscoff, CS 90074, F-29688, Roscoff cedex, France.
authors.forename: Gurvan
authors.initials: G
authors.lastname: Michel
authors.affiliation: Institute for Evolution and Biodiversity, Westfälische Wilhelms-University of Münster, Hüfferstrasse 1, D-48149 Münster, Germany.|Institute for Computer Science, Heinrich Heine University, D-40255 Duesseldorf, Germany.
authors.forename: Anna
authors.initials: A
authors.lastname: Kersting
authors.affiliation: Stazione Zoologica Anton Dohrn, Villa Comunale, 80121 Naples, Italy.
authors.forename: Chiara
authors.initials: C
authors.lastname: Lauritano
authors.affiliation: Department of Plant Systems Biology, VIB and Department of Plant Biotechnology and Bioinformatics, Ghent University, Technologiepark 927, B-9052 Ghent, Belgium.
authors.forename: Rolf
authors.initials: R
authors.lastname: Lohaus
authors.affiliation: Department of Biological and Environmental Sciences, Bioinformatics Infrastructure for Life Sciences (BILS), University of Gothenburg, Medicinaregatan 18A, 40530 Gothenburg, Sweden.
authors.forename: Mats
authors.initials: M
authors.lastname: Töpel
authors.affiliation: Sorbonne Université, UPMC Univ Paris 06, CNRS, UMR 8227, Integrative Biology of Marine Models, Station Biologique de Roscoff, CS 90074, F-29688, Roscoff cedex, France.
authors.forename: Thierry
authors.initials: T
authors.lastname: Tonon
authors.affiliation: Department of Plant Systems Biology, VIB and Department of Plant Biotechnology and Bioinformatics, Ghent University, Technologiepark 927, B-9052 Ghent, Belgium.
authors.forename: Kevin
authors.initials: K
authors.lastname: Vanneste
authors.affiliation: Department of Energy Joint Genome Institute, 2800 Mitchell Dr., #100, Walnut Creek, California 94598, USA.
authors.forename: Mojgan
authors.initials: M
authors.lastname: Amirebrahimi
authors.affiliation: GEOMAR Helmholtz Centre for Ocean Research-Kiel, Evolutionary Ecology, Düsternbrooker Weg 20, D-24105 Kiel, Germany.
authors.forename: Janina
authors.initials: J
authors.lastname: Brakel
authors.affiliation: Environmental and Marine Biology, Faculty of Science and Engineering, Åbo Akademi University, Artillerigatan 6, FI-20520 Turku/Åbo, Finland.
authors.forename: Christoffer
authors.initials: C
authors.lastname: Boström
authors.affiliation: Department of Energy Joint Genome Institute, 2800 Mitchell Dr., #100, Walnut Creek, California 94598, USA.
authors.forename: Mansi
authors.initials: M
authors.lastname: Chovatia
authors.affiliation: Department of Energy Joint Genome Institute, 2800 Mitchell Dr., #100, Walnut Creek, California 94598, USA.|HudsonAlpha Institute for Biotechnology, 601 Genome Way NW, Huntsville, Alabama 35806, USA.
authors.forename: Jane
authors.initials: J
authors.lastname: Grimwood
authors.affiliation: Department of Energy Joint Genome Institute, 2800 Mitchell Dr., #100, Walnut Creek, California 94598, USA.|HudsonAlpha Institute for Biotechnology, 601 Genome Way NW, Huntsville, Alabama 35806, USA.
authors.forename: Jerry W
authors.initials: JW
authors.lastname: Jenkins
authors.affiliation: Marine Ecology Group, Nord University, Postbox 1490, 8049 Bodø, Norway.
authors.forename: Alexander
authors.initials: A
authors.lastname: Jueterbock
authors.affiliation: Amplicon Express, 2345 NE Hopkins Ct., Pullman, Washington 99163, USA.
authors.forename: Amy
authors.initials: A
authors.lastname: Mraz
authors.affiliation: Groningen Institute of Evolutionary Life Sciences (GELIFES), University of Groningen, PO Box 11103, 9700 CC Groningen, The Netherlands.
authors.forename: Wytze T
authors.initials: WT
authors.lastname: Stam
authors.affiliation: Department of Energy Joint Genome Institute, 2800 Mitchell Dr., #100, Walnut Creek, California 94598, USA.
authors.forename: Hope
authors.initials: H
authors.lastname: Tice
authors.affiliation: Institute for Evolution and Biodiversity, Westfälische Wilhelms-University of Münster, Hüfferstrasse 1, D-48149 Münster, Germany.
authors.forename: Erich
authors.initials: E
authors.lastname: Bornberg-Bauer
authors.affiliation: School of Marine Science and Policy, Department of Plant and Soil Sciences, Delaware Biotechnology Institute, University of Delaware, 15-Innovation Way, Newark, Delaware 19711, USA.
authors.forename: Pamela J
authors.initials: PJ
authors.lastname: Green
authors.affiliation: Marine Ecology and Evolution, Centre for Marine Sciences (CCMAR), University of Algarve, 8005-139 Faro, Portugal.
authors.forename: Gareth A
authors.initials: GA
authors.lastname: Pearson
authors.affiliation: Stazione Zoologica Anton Dohrn, Villa Comunale, 80121 Naples, Italy.
authors.forename: Gabriele
authors.initials: G
authors.lastname: Procaccini
authors.affiliation: King Abdullah University of Science and Technology (KAUST), Red Sea Research Center (RSRC), Thuwal 23955-6900, Saudi Arabia.
authors.forename: Carlos M
authors.initials: CM
authors.lastname: Duarte
authors.affiliation: Department of Energy Joint Genome Institute, 2800 Mitchell Dr., #100, Walnut Creek, California 94598, USA.|HudsonAlpha Institute for Biotechnology, 601 Genome Way NW, Huntsville, Alabama 35806, USA.
authors.forename: Jeremy
authors.initials: J
authors.lastname: Schmutz
authors.affiliation: GEOMAR Helmholtz Centre for Ocean Research-Kiel, Evolutionary Ecology, Düsternbrooker Weg 20, D-24105 Kiel, Germany.|University of Kiel, Faculty of Mathematics and Natural Sciences, Christian-Albrechts-Platz 4, 24118 Kiel, Germany.
authors.forename: Thorsten B H
authors.initials: TB
authors.lastname: Reusch
authors.affiliation: Department of Plant Systems Biology, VIB and Department of Plant Biotechnology and Bioinformatics, Ghent University, Technologiepark 927, B-9052 Ghent, Belgium.|Genomics Research Institute, University of Pretoria, Hatfield Campus, Pretoria 0028, South Africa.|Bioinformatics Institute Ghent, Ghent University, Ghent B-9000, Belgium.
authors.forename: Yves
authors.initials: Y
authors.lastname: Van de Peer
mesh_terms: D000064:Acclimatization / Q000235:genetics; D000222:Adaptation, Physiological / Q000235:genetics*; D002473:Cell Wall / Q000737:chemistry; D005030:Ethylenes / Q000096:biosynthesis; D019143:Evolution, Molecular; D020440:Gene Duplication; D017343:Genes, Plant / Q000235:genetics; D018745:Genome, Plant / Q000235:genetics*; D053858:Metabolic Networks and Pathways; D008969:Molecular Sequence Data; D009792:Oceans and Seas; D064587:Osmoregulation / Q000235:genetics; D010802:Phylogeny; D018515:Plant Leaves / Q000378:metabolism; D054046:Plant Stomata / Q000235:genetics; D011058:Pollen / Q000378:metabolism; D054712:Salinity; D055049:Salt Tolerance / Q000235:genetics; D012623:Seawater; D012624:Seaweed / Q000235:genetics; D013729:Terpenes / Q000378:metabolism; D035962:Zosteraceae / Q000235:genetics*
chemical_list: D005030:Ethylenes; D013729:Terpenes; C036216:ethylene