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Phlorotannins As UV-Protective Substances in Early Developmental Stages of Brown Algae

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Phlorotannins As UV-Protective Substances in Early Developmental Stages of Brown Algae 651 2012 Phlorotannins as UV-protective Substances in early developmental Stages of Brown Algae Phlorotannine als UV-Schutzsubstanzen in frühen Entwicklungsstadien von Braunalgen Franciska S. Steinhoff ALFRED-WEGENER-INSTITUT FÜR POLAR- UND MEERESFORSCHUNG in der Helmholtz-Gemeinschaft D-27570 BREMERHAVEN Bundesrepublik Deutschland ISSN 1866-3192 Hinweis Notice Die Berichte zur Polar- und Meeresforschung The Reports on Polar and Marine Research are werden vom Alfred-Wegener-Institut für Polar- issued by the Alfred Wegener Institute for Polar und Meeresforschung in Bremerhaven* in un- and Marine Research in Bremerhaven*, Federal regelmäßiger Abfolge herausgegeben. Republic of Germany. They are published in irregular intervals. Sie enthalten Beschreibungen und Ergebnisse They contain descriptions and results of der vom Institut (AWI) oder mit seiner Unter- investigations in polar regions and in the seas stützung durchgeführten Forschungsarbeiten in either conducted by the Institute (AWI) or with its den Polargebieten und in den Meeren. support. Es werden veröffentlicht: The following items are published: — Expeditionsberichte — expedition reports (inkl. Stationslisten und Routenkarten) (incl. station lists and route maps) — Expeditions- und Forschungsergebnisse — expedition and research results (inkl. Dissertationen) (incl. Ph.D. theses) — wissenschaftliche Berichte der — scientific reports of research stations Forschungsstationen des AWI operated by the AWI — Berichte wissenschaftlicher Tagungen — reports on scientific meetings Die Beiträge geben nicht notwendigerweise die The papers contained in the Reports do not Auffassung des Instituts wieder. necessarily reflect the opinion of the Institute. The „Berichte zur Polar- und Meeresforschung” continue the former „Berichte zur Polarforschung” * Anschrift / Address Editor: Alfred-Wegener-Institut Dr. Horst Bornemann für Polar- und Meeresforschung D-27570 Bremerhaven Assistant editor: Germany Birgit Chiaventone www.awi.de Die "Berichte zur Polar- und Meeresforschung" (ISSN 1866-3192) werden ab 2008 als Open-Access-Publikation herausgegeben (URL: http://epic.awi.de). Since 2008 the "Reports on Polar and Marine Research" (ISSN 1866-3192) are available as open-access publications (URL: http://epic.awi.de) Phlorotannins as UV-protective Substances in early developmental Stages of Brown Algae Phlorotannine als UV-Schutzsubstanzen in frühen Entwicklungsstadien von Braunalgen Franciska S. Steinhoff Please cite or link this publication using the identifier hdl:10013/epic.40016 or http://hdl.handle.net/10013/epic.40016 ISSN 1866-3192 Erkennen heißt nicht zerlegen, auch nicht erklären. Es heißt Zugang zur Schau finden. Aber um zu schauen, muss man erst teilnehmen. Das ist eine harte Lehre. Antoine de Saint-Exupéry Franciska S. Steinhoff Norwegian University of Science and Technology (NTNU) NO-7491 Trondheim Norway [email protected] Die vorliegende Arbeit ist die inhaltlich geringfügig veränderte Fassung einer Dissertation, die am Alfred-Wegener-Institut in der Sektion Funktionelle Ökologie des Fachbereichs Biowissenschaften, im Fachbereich Marine Biologie an der Universität Bremen sowie und an der internationalen Bremer Graduiertenschule für marine Wissenschaften (GLOMAR) angefertigt und 2010 dem Fachbereich 2 für Biologie und Chemie der Universität Bremen vorgelegt wurde. This publication is a slightly altered version of a cumulative dissertation conducted at the Alfred-Wegener-Institute for Polar and Marine Research, at the University of Bremen and at the Bremen International Graduate School for marine Sciences (GLOMAR) and was submitted 2010 to the Department of Biology and Chemistry at the University of Bremen. CONTENTS LIST OF ABBREVIATIONS…………………………………………………………... III SUMMARY……………………………………………………………………………... V ZUSAMMENFASSUNG…..…………………………………………………………… VIII LIST OF PUBLICATIONS……………………………………………………………. XII 1 Introduction…………………………………………………………………………... 1 1.1. Seaweeds……………………………………………………………………………... 1 1.1.1. Seaweed communities in the Arctic……………………………………….. 2 1.1.2. Developmental life-cycle of kelps………………………………………… 3 1.1.3. Ultrastructure of kelp zoospores…………………………………………... 4 1.2. Stratospheric ozone layer, greenhouse effect and variability of UV radiation……... 4 1.2.1. Stratospheric ozone layer, UV radiation and PAR……………………….... 5 1.2.2. Impact of UV radiation and high PAR irradiances on algae..…………….. 6 1.3. Climate change and enhanced temperatures in the Arctic……………………...…… 7 1.4. Protective, stress and storage compounds in zoospores……………..…………….... 8 1.4.1. Phlorotannins……………………………………………………………… 8 1.4.2. Reactive oxygen species…………………………………………….…….. 10 1.4.3. Fatty acids in kelps………………………………………………………… 10 1.5 Aims of the thesis…………………………………………….…………………..….. 11 2 Methodological considerations……………………………………………………… 12 2.1. The Kongsfjord, an Arctic Fjord System…………………………………………… 12 2.1.1. The physical environment of Kongsfjorden……………………………… 13 2.1.1.1. Water temperature……………………………………………… 13 2.1.1.2. Underwater radiation regime…………………………………... 13 2.1.2. Study area and sample collection………………………………………... 14 2.2. Experimental design, set-up and investigated parameters……………………….… 14 2.2.1. Underwater simulations ……………………………………………….... 16 2.3. Experimental design in relation to statistics……………………………………….. 17 2.4. Radiation treatments and measurements…………………………………………… 17 2.5. Germination as a fitness parameter………………………………………………… 18 2.6. Ultrastructructure of gametophytes and reactive oxygen species (ROS)………….. 19 2.7. Quantitative and qualitative analysis of fatty acids………………………………… 19 2.8. Phlorotannin extraction…………………………………………………………….. 20 2.9. Quantitative phlorotannin determination…………………………………………… 21 3 Summary of Results………………………………………………………………….. 23 3.1. UV radiation at Ny-Ålesund and in Kongsfjorden…………………………………. 23 3.2. Effects of a changing environment on spores………………………………………. 24 3.2.1. Germination of various brown algal zoospores…………………………... 24 3.2.2. Phlorotannin content in zoospores and their surrounding medium………. 25 I 3.2.2.1. Phlorotannin variability over time……………………………… 25 3.2.2.2. Phlorotannin variability under UV radiation and in various temperatures…………………………………………………..… 26 3.2.2.3. Phlorotannin content and vertical zonation pattern of algae…… 28 3.2.3 Fatty acid composition in kelp zoospores and juvenile gametophytes……. 29 3.2.3.1. Effects of time, UV radiation and low vs high PAR on fatty acid content and composition……………………………………. 30 3.2.3.2. Coherence of fatty acids and phlorotannins…………………….. 31 3.2.4. Microscopical observations of UV radiation and temperature effects……. 32 4 Synoptic Discussion…………………………………………….…………………….. 34 4.1. Experimental radiation treatments and measurements……………………………… 34 4.2. Zoospores, a challenging developmental stage?......................................................... 35 4.3. The notional role of phlorotannins in brown macroalgae…………………………... 36 4.3.1. Phlorotannins, induction in relation to season and species?........................ 36 4.3.2. Phlorotannin expression in relation to habitat and age of the algae?.......... 37 4.3.3. Phlorotannin expression in relation to radiation tolerance and exposure time, ROS and vertical zonation?................................................................ 39 4.3.4. Phlorotannins - protective substances for recruits?..................................... 42 4.4. Fatty acid composition, variable or reliable metabolic parameter?............................ 42 4.4.1. Fatty acid composition in relation to species……………………………... 43 4.4.2. Fatty acid composition in relation to UV radiation or/and high PAR.…… 44 4.5. Short- and mid-term radiation and temperature effects on brown algal propagules.. 45 4.6. Conclusion………………………………………………………………………….. 47 4.7. Future perspectives…………………………………………………………………. 49 References………………………………………………………………………………. 51 Original Publications…………………………………………………………………... 66 Publication I……………………………………………………………………………. 67 Publication II…………………………………………………………………………… 76 Publication III………………………………………………………………………….. 78 Publication IV………………………………………………………………………….. 80 Publication V…………………………………………………………………………… 93 Publication VI………………………………………………………………………….. 110 Acknowledgements…………………………………………………………………….. 121 II List of Abbreviations e.g. Exampli gratia (for example) % Percent ˜ Approximately EM Electron microscopy < Smaller than FA Fatty acid > Greater than FC Folin-Ciocalteu ° Degree FD Folin-Denis °C Degree Celsius Feb February µ Micro GC Gas chromatography µg Microgram h Hours µm Micrometer H2O2 Hydrogen peroxide µmol Micromole HPLC High performance liquid chromatography Acetyl-CoA Acetyl-Co-enzyme A IR Infrared Apr April Jan January Aug August Jul July C Carbon Jun June CIE Commission Internationale de l’Éclairage kDa Kilo Dalton CPD Cyclobutane pyrimidine kg Kilogram dimer km Kilometer d Days L Lipid globules Da Dalton LM Light microscopy Dec December m Meter DNA Desoxyribonucleic acid MAA Mycosporine-like amino DOC Dissolved organic matter acid DU Dobson unit Mar March mol Mole E East mRNA Messenger Ribonucleic acid III Abbreviations MUFA Monosaturated fatty acids UV-A Ultraviolet radiation A (320-400 nm) N North UV-B Ultraviolet radiation B n Number of replicates (280-320 nm) NBT Nitro-blue tetrazolium UV-C Ultraviolet radiation C (220-280 nm) nm Nanometer UVR Ultraviolet radiation Nov November (220-400 nm) Oct October W Watt [SI Unit] OH Hydroxyl WSC West Spitsbergen Current p Probability λ Wavelength [nm] PA Photosynthetically active radiation (PAR) + UV- A(320-700
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