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Green Algae in Marine Phytoplankton

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Green Algae in Marine Phytoplankton Diversity of marine phytoplankton Images de Plankton@net Nathalie Simon January 2012 – UE EPHYBIO – Master 2 Phytoplankton : « wandering plants » mg/m3 0.1 1% of Chl 0.5 on earth 10 45% of Net Primary Production Ch a in Oceans Biological pump Enrichment in inorganic C Ci enrichment maintains excess C02 in oceans compared to atmosphere (Ocean : 300 ppm more CO2) …plancton Micro- Méso- (20-200 µm) (200µm-2mm) Qqs mm Nano- (2-20 µm) Pico- (0,2-2 µm) 0,4 µm Finlay, 2002 Cyanobacteria Cyanobacteria -Unicellular, trichoms, filaments -Bacterial Cellular Envelopp PE I -Thylacoïds concentric, Chl a, carotenes, APC PC phycobilisomes (except for few genera PE II such as Prochlorococcus) -Genetic material: circular DNA, no histones -Possibility for N2 fixation (hétérocystes) -Gaz Vacuoles (plankton) Synechocystis glycogène granule de cyanophycine carboxysome ribosome Mb externe 100 nm Peptidoglucane Liberton et al., 2006 Mb plasmique Cyanobacteria : gram- Bacteria Marine planktonic cyanobacteria: ~14 genera? Trichodesmium -Trichomes organised in filaments -Blooms (tropics) -atm N2 Fixation Richelia intracellularis ©LOB/K. Leblanc -Trichomes with heterocysts(Fixation N2 atm.) - Symbiosis with diatoms (Guinardia, Hemiaulus,…) ©LOB/K. Leblanc (Photo by Dave Caron, Woods Hole Oceanographic Institution) Synechococcus Prochlorococcus « Crocosphaera » Unicellular Several genetic Several genetic Low genetic diversity clades clades Antenna with Antenna with Antenna with Divinyl Chla/b phycobilisomes phycobilisomes No N2 fixation No N2 fixation N2 Fixation 2 functionnal groups 1. Non diazotroph picocyanobacteria : major contribution to primary production in oligotrophic ocean (ex. adaptations : reduced size) 2. Filamentous diazotrophs : More than ½ of N import in tropics. Primitive organisms with characters similar to those of PS Cyanobacteria p (-2,7 Billion years) are at the origin of atmospheric oxygenation (-2,3 Billion years). 2 functionnal groups 1. Non diazotroph picocyanobacteria : major contribution to primary production in oligotrophic ocean (ex. adaptations : reduced size) 2. Filamentous diazotrophs : More than ½ of N import in tropics. Primitive organisms with characters similar to those of PS Cyanobacteria p (-2,7 Ma y) are at the origin of atmospheric oxygenation (-2,3 Ma ans), and are at the origin of all plastids. Plastids form a Synechococcus Prochlorococcus monophyletic group Phormidium Prochlorothrix Leptolyngbya Chamaesiphon Nostoc, Calothrix, Scytonema, Cyano- Anabaena, Cylindrospermum Bacteria Oscillatoria, Trichodesmium, + Microcoleus, Arthrospira, Lyngbya Plastids Gloeothece, Gloeocapsa, Synechocystis Microcystis, Pleurocapsa, Prochloron Plastids Pseudanabaena Limnothrix Unicellulars thermophiles Strains phylogenetically close to Gloeobacter rDNA 16S Gloeobacter Turner (1997, 1999) Photosynthesis acquired by eukaryotes through endosymbiosis Primary endosymbiosis Eukaryotes consensus phylogeny SAR Plantae Habrobia Unikonts Excavates Bauldauf 2008 Reviers 2011 Diatoms, dinoflagellates and haptophytes (coccolithophores) 3 major groups of the micro- and nano-phytoplankton Diatoms Belong to Heterokonta (Stramenopiles) Phylum Ochrophyta Lee (2008) 40 % Of net Primary production by phytoplankton 40% of described species within phytoplankton 50 % of exported C Microplankton (nanoplankton) Chap 11 vidéo Generalities about diatoms -Photosynthetic, unicellular (forming colonies) -Frustule = silicified cell wall -Brown plastids -10 000 species (> 2000 marine planktonic -Ecologically diverse -Fossils and geological formations “diatomite” “tripoli” ou “kieselguhr” Frustule Epitheca Hypotheca Epivalve Cingular band Source : wikipedia Drawings from : W. Kooistra Le frustule Vues valvaires Vue cingulaire Les processus : projections à parois silicifiées Thalassiosira eccentrica Plankton@net Ornementations Processus (projections de silice) Pores Areolae Chloroplast : Envelop with 4 membranes Groups of 3 thylacoïdes, lamella, pyrénoïd(s) or not Pigments : Chl a, c, fucoxanthin + caroténoïds Storage material: Chrysolaminarin polyphosphates Lipid droplets Van den Hoek, 1995 Origin of diatoms plastids 2 complete genomes E I = Cyanobacteria Moustafa et al. Science 2010 : Origin of diatoms EII= Red algea genes identified through phylogenomics (among genes identified as originating from green or red algae) Plastid originates from a Iiary endosymbiosis with red algae A Iiary endosymbiosis with a green algae could have left traces??? Radial Centrics Corethron Stephanopyxis Coscinodiscus Diapositive : W. Kooistra Multi-polar Centrics Odontella Thalassiosira Chaetoceros Diapositive : W. Kooistra Araphid Pennates Fragilaria Synedra Licmophora Asterionella Diapositive : W. Kooistra Raphid Pennates Pseudo-nitzschia Entomoneis Campylodiscus Diapositive : W. Kooistra Radial centrics Multi-polar centrics Araphid pennates Raphid pennates SSU 0.01 Diapositive : W. Kooistra 0 Frustules fossiles T Presence of pennates with raphe 65 90 Presence of pennates without raphe K Antarctic ODP-Leg 693 Gersonde and Harwood, 1990 Australia Nikolaev and Harwood, 1997 120 Radial centrics and few multipolar 145 Calyptosporium, (Korea) Harwood et al., in prep. J Radial centrics Pyxidicula, (Germany) Rothpletz, 1896 180 Radial centrics 207 Tr De W. Kooistra modifiée Ancestral benthic life style Planktonic diatoms appeared several times in evolution Benthic epiphytic Few pennate planktonic diatoms Mobile on substrate 0.01 Modified from : W. Kooistra Chaetoceros et al. Chaetoceros Eucampia Adaptations to live in the plankton © Jan Rines Diapositive : W. Kooistra Thalassiosirales Thalassiosira Skeletonema Adaptations to live in the plankton Diapositive : W. Kooistra Asterionellopsis Thalassionema Asterionellopsis Asterionella Thalassionema Asterionella Pseudo-nitzschia and Fragilariopsis Diapositive : W. Kooistra Trades-off : selection /adaptation Many opportinistic « bloomers » Frustule • Protection against predation • Flottability Large vacuole •Stocks of NO3-, PO42- •Allows to lower the amount available to competitors. Dinoflagellates Alveolata, Dinophyta Dinophyceae Important part of net primary production in the oceans 40% of marine phytoplankton species described Nano - Microplankton -Unicellular (may form colonies) -Cell wall : Cellulose in alveoles -2 flagella (transverse + Epithèque longitidinal) Epicône -Brown, red, green plastids -Taxonomic diversity: 5000 species (2000 planktonic photosynthetic) D G Hypothèque, hypocône Vue ventrale Alveoles with or without cellulose plaques Naked / armored Karenia brevis Alexandrium tamarense Dinophysis acuta From Lee (1989) and plankton@net Disposition « desmocontée » Dinokont disposition 2 flagelles à l’apex Flagella inserted ventraly, in silts Transverse Fl. (in cingulum) Longitudinal Fl. (in sulcus) Karenia brevis Peridinium cinctum Dorsal vue Ventral vue Modified from Lee, 1989 Ultrastructure Trichocyst Chloroplasts – most frequent Envelopp with 3 membranes, thylacoïds by 3 Chl a, c, peridinin Nucleus (condensed chromosomes - Histones very peculiar - ~100 x more DNA than other euks) Phycologia, mai 2004 Main storage: starch Le dinokaryon, un noyau très particulier Interphase Début division Ségrégation des chromosomes (avec mise en place d’un canal cytoplasmique) Division du noyau B : Corps basaux, K : kinétochores, C : Chromosomes, NM : membrane nucléaire, Mt : Microtubules RCC1488 : Lepidodinium chlorophorum (isolée par I. Probert en Manche) Photos de Chantal Billard Lepidodinium chlorophorum Prasinophyceae : Pyramimonas Pyramimonas SEM of the flagellated prasinophyte Pyramimonas gelidicola Image: Sandy Melloy Moestrup et Walne J. Cell Sci. 36, 437-459 0979) Gould et al., 2008 Dinoconts Epitheca Epicone Hypotheca, hypocone Desmoconts (Prorocentrales) D ’après Lee, 1989 Gymnodiniales No theca (or thin) Cingulum Suessiales Péridiniales Thick theca Cingulum Gonyaulacales 2 flagella at apex Theca with elaborated extensions Prorocentrales Theca divided in 2 parts Dynophysiales No cigulum Adaptation to planktonic life • Colonies, cell wall spines and • Ethology : nyctemeral migrations extensions Profil de Chl a durant un bloom de dinoflagellés (d ’après Eppley et al. 1968) Haptophytes incl. coccolithophores - 10 % of described marine phytoplankton spcecies (380 esp.) - Major role in biogeochemical cycles (C, du S) - Nanoplankton (picoplankton) Organic scales with/without calcification Emiliania huxleyi Chrysochromulina Coccolithus Hymenomonas http://microscope.mbl.edu/baypaul/microscope/general /page_01.htm Organic microfibrillous scales •Proximal face : radially organised, sectors Chrysochromulina / Prymnesium •Distal face : variable, may include projections •May serve as a matrix for biomineralisation (coccolithophores) Phaeocystis Cosmopolitan Harmful blooms (North Sea) Références : Plankton@net, EOL, Lee (2008) 2 µm 0,2 µm 2 µm Zingone et al. 2011) – Phaeocystis antarctica Coccoliths • Organic scales may be • 2 major types used as a matrix for biomineralisation 1/ Hétérococcoliths Intact Coccolith Radial arrangment of calcite cristals in intracellular vacuoles 2/ Holococcolithes Scale « matrix » Extracellular precipitation of small hexahedric CaCO3 cristals Dissolved coccolith Emiliania huxleyi 2n n Organic scales Coccoliths Image: Remote Sensing Data Analysis Service (RSDAS) www.npm.ac.uk/rsdas/ of the Plymouth Marine Laboratory (PML) The haptonema • Variable size • 3 concentric membranes, 7 microtubules • Mouvment and capture of preys Chrysochromulina Lee (2008) Caractères cytologiques et ultrastructuraux 1 Haptonème (6 à
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