Aquatic ecology BIO 4400 Phytoplankton and primary production-3 Photo: Jahn Throndsen Bente Edvardsen 2009 Content • Algal phylogeny and evolution • Marine phytoplankton groups • • • • • Diatoms (Kiselalger) Dinoflagellates (Fureflagellater) Haptophytes (Svepeflagellater) Cyanobacteria (Cyanobakterier) Other groups Eukaryotic tree of life Baldauf 2003. Science 300: 1703-6 From Baldauf 2003. Science 300: 1703-6. Modified by N. Simon Origin of plastids in algae: The endosymbiosis theory PROKARYOTA Division CYANOPHYTA (cyanobakterier eller blågrønnalger) 2000 species class Cyanophyceae EUKARYOTA Division GLAUCOPHYTA 14 species Division RHODOPHYTA (rødalger) 5000 species class Cyanidiophyceae class Bangiophyceae class Florideophyceae Division HETEROKONTOPHYTA (heterokonte alger) class Chrysophyceae (Gullalger) ca 1000 species class Chrysomerophyceae class Bolidophyceae class Pelagophyceae class Pinguiophyceae class Xanthophyceae class Phaeothamniophyceae class Eustigmatophyceae class Bacillariophyceae (kiselalger eller diatomeer) 12 000 - ? species class Raphidophyceae (nålflagellater) 25 species class Dictyochophyceae (kiselflagellater) 40 species class Phaeophyceae (brunalger) 1800 species Algal taxonomy Algae can be divided into 10 divisions Division HAPTOPHYTA (svepeflagellater) 300 species class Prymnesiophyceae class Pavlovophyceae Division CRYPTOPHYTA (svelgflagellater) 126 species Important classes in marine plankton are in green class Cryptophyceae Division DINOPHYTA (fureflagellater eller dinoflagellater) ca 2000 species class Dinophyceae Division EUGLENOPHYTA (øyealger) 800 species class Euglenophyceae Division CHLORARACHNIOPHYTA 6 species Division CHLOROPHYTA (grønnalger sensu lato) class Prasinophyceae (olivengrønnalger) 200 species class Chlorophyceae 1000 species class Ulvophyceae 950 species class Trebouxiophyceae ca. 15 genera class Charophyceae ca 5000 species From Throndsen, Rueness and Fredriksen, BIO 1200 Division Heterokontophyta Class Bacillariophyceae Diatoms (Kiselalger) Characteristics (kjennetegn) • silicate cell wall (kiselskall) • autotrophic • dominate the vernal bloom (våroppblomstringen) • Planktonic froms are often chain-forming • diplonts Diatoms Kiselalger Morphology • Cell wall (frustule) of silicate (as glass) (SiO2)n - (H2O)n • In two parts: box (hypotheca) + lid (epitheca) pennate symmetry along a line centric symmetry around a point Centric diatoms Thalassiosira Chaetoceros Guinardia Rhizosolenia Odontella Fra N. Simon, SBR Pennate diatoms Thalassionema Navicula Nitzschia Fra N. Simon, SBR Cell wall (cellevegg) valva Girdle (Bånd) JT04 Cross section (tverrsnitt) of cell wall skallside skallkappe beltebånd Cell wall = frustule from valve face (fra skallsiden) from girdle face (fra båndsiden) Thalassiosira eccentrica Axe pervalvaire Fra N. Simon, SBR SEM Areols : openings in the frustule MO SEM Processes: tube formed structures of silicate Raphe (rafe): a canal in some pennate diatoms JT Colony formation In Skeletonema the silicate processes are fused Labiate processes JT Photo by G.R. Hasle colonies Chaetoceros JT raphe • • • • simple raphe canal raphe central nodule pseudoraphe Round & Crawford 1990 Diploneis sp. raphe Central node (sentralknute) SEM-graph from ”Norsk kystplanktonflora” JT Diatoms – cell division Division Dinophyta Class Dinophyceae Dinoflagellates (Fureflagellater) Characteristics (Kjennetegn) • Transvers and longitudinal flagella • Large nucleus – dinokaryon • Cell wall of cellulose (plates) in some • Auto- and heterotrophy • Haplonts Dinoflagellates (Fureflagellater) Ventral side right left JT cell wall Amphiesma is a vacuole system just beneath the cell membrane that form cellulose plates in thecate, but not athecate Peridinium cinctum Dorsal side Ventral side flagella and groves (furer) TVERRFURE CINGULUM LENGDEFURE SULCUS JT two flagella in two groves Transverse flagellum (tverrflagell) Karenia brevis streng longitudinal flagellum (lengdeflagell) Winged transverse flagellum (Vinget tverrflagell) dinokaryon dinokaryon Large nucleus with condensed chromosomes Pentapharsodinium dalei Main morphotypes of dinoflagellates prorocentroid Prorocentrum dinophysoid Dinophysis peridinoid Protoperidinium gymnodinoid Karenia JT Ceratium morphology Ceratium tripos SEM-graph JT Ceratium furca dinoflagellates - nutrition Phagotrophy with pallium in Protoperidinium spinulosum dinoflagellates - nutrition Phagotrophy with peduncle in Stylodinium Oedogonium dinoflagellates - nutrition Protoperidinium depressum Oil droplets JT dinoflagellates - nutrition Dinophysis acuta eats M. rubra Myrionecta rubra is a ciliate that eats cryptophytes dinoflagellates – life cycle Seasonal cycle with cyst formation Lingulodinium polyedrum dinoflagellates – life cycle Resting stage in dinoflagellates - cysts Lingulodinium polyedrum (cyste eller hvilespore) Bioluminescence (morild) Noctiluca scintillans Division Haptophyta (Svepeflagellater) Flagellates, most nanoplanktonic (usually 4-20 µm) Prymnesiophyceae (Pleurochrysis ) Pavlovophyceae (Pavlova ) JT04 Haptophyta General features haptonema • Two smooth flagella • Haptonema • Cells covered by organic scales flagella chloroplast • Calcified scales (coccoliths) may be present • Heteromorphic haplontdiplont life cycle Throndsen scales Flagella and haptonema Haptonema that can coil or bend haptonema flagellum JT04 Class Prymnesiophyceae Class Pavlovophyceae JT04 Nutrition in haptophytes mixotrophy Particle handling JT04 Division Haptophyta (Svepeflagellater ) Including coccolithophores (kalkflagellater) Chrysochromulina Coccolithus Emiliania huxleyi Discosphaera tubifera Hymenomonas 300 species, Almost all are marine Haptophytes have species specific scales seen in the electron microscope 5 µm Coccolithophores have calcified coccoliths 5 µm 1 µm 1 µm Organic scales Blooms of coccolithophores can be observed from space by satellites when light is reflected by the coccoliths Colony formation Phaeocystis spp. P. pouchetii P. globosa Some haptophyte blooms may be toxic to fish Chrysochromulina polylepis “Algekatastrofen”, Skagerrak May 1988 Prymnesium parvum in Texas Cyanobacteria (Division Cyanophyta) important picoplankton Prochlorococcus DIVISION Heterokontophyta = Ochrophyta = Stramenopila Chrysolepidomonas (Chrysophyceae) Chattonella (Raphidophyceae) 13 classes Laminaria (Phaeophyceae) Thalassiosira (Bacillariophyceae) Aureoumbra (Pelagophyceae) Mallomonas (Synurophyceae) From N. Simon, Roscoff Class Dictyochophyceae (kiselflagellater) Dictyocha speculum Photo: Jahn Throndsen The dictyochophyte Pseudochattonella farcimen - an introduced species? • Pseudochattonella had not been reported from European waters before 1993. • Pseudochattonella verruculosa has produced fish-killing blooms in Japan