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Characterization of the Life-Cycle Stages of the Coccolithophore Emiliania Huxleyi and Their Responses to Ocean Acidification

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Characterization of the Life-Cycle Stages of the Coccolithophore Emiliania Huxleyi and Their Responses to Ocean Acidification Characterization of the life-cycle stages of the coccolithophore Emiliania huxleyi and their responses to Ocean Acidification Dissertation zur Erlangung des akademischen Grades eines Doktors der Naturwissenschaften - Dr. rer. nat. - Am Fachbereich 2 (Biologie/Chemie) der Universität Bremen Vorgelegt von Sebastian Rokitta im September 2012 I II Sebastian Rokitta, Dipl. Biol. Alfred-Wegener-Institut für Polar- und Meeresforschung in der Helmholtz-Gemeinschaft ERC Nachwuchsgruppe PhytoChange / Marine Biogeosciences Am Handelshafen 12, 27570 Bremerhaven, Germany [email protected] III IV Table of Contents Table of Contents ............................................................................................................................................. 1 PREFACE ........................................................................................................................................................... 3 Disclaimer ........................................................................................................................................... 3 Acknowledgements ............................................................................................................................ 5 Summary ............................................................................................................................................ 7 Zusammenfassung ............................................................................................................................. 8 List of abbreviations ......................................................................................................................... 11 List of figures .................................................................................................................................... 12 INTRODUCTION .............................................................................................................................................. 13 Global change and the Anthropocene .............................................................................................. 13 The human carbon perturbation ...................................................................................................... 14 Seawater carbonate chemistry ........................................................................................................ 15 Ocean Acidification........................................................................................................................... 18 Phytoplankton-driven carbon pumps ............................................................................................... 20 Evolutionary history of coccolithophores ......................................................................................... 22 The Emiliania huxleyi morphospecies complex ................................................................................ 25 The dualism of biomass and calcite production in diploid E. huxleyi ............................................... 27 Haplo-diplontic life cycling in E. huxleyi ........................................................................................... 29 Aims of this thesis ............................................................................................................................. 30 PUBLICATIONS ............................................................................................................................................... 33 List of publications ............................................................................................................................ 33 The first author´s contribution to the publications .......................................................................... 33 Publication I: Transcriptome analyses reveal differential gene expression patterns between life- cycle stages of Emiliania huxleyi (Haptophyta) and reflect specialization to different ecological niches ................................................................................................................................................ 35 Publication II: Effects of CO2 and their modulation by light in the life-cycle stages of the coccolithophore Emiliania huxleyi .................................................................................................... 47 Publication III: Ocean Acidification affects carbon allocation and ion-homeostasis at the transcriptomic level in the life-cycle stages of Emiliania huxleyi ..................................................... 61 DISCUSSION ................................................................................................................................................... 73 Major findings for the diploid life cycle stage .................................................................................. 73 1 Major findings for the haploid life cycle stage ................................................................................. 74 Emerging response patterns and unifying physiological concepts .................................................. 75 Perspectives and research outlook ................................................................................................... 80 CONCLUSION .................................................................................................................................................. 83 REFERENCES ................................................................................................................................................... 85 APPENDIX....................................................................................................................................................... 97 Publication: Implications of observed inconsistencies in carbonate chemistry measurements for ocean acidification studies ............................................................................................................... 97 Bookchapter: Mechanism of coccolith synthesis ............................................................................ 104 Publication: Emiliania´s pan genome drives the phytoplankton´s global distribution ................... 106 2 PREFACE Disclaimer I hereby assure that I myself have performed the work described in this doctoral thesis. All external resources used as well as the contributions made to the publications are appropriately stated. _______________________________ 3 4 Acknowledgements I am exceptionally grateful to Björn Rost. Besides for your guidance, I thank you for your contagious curiosity that made me embrace ever new and exciting topics. Thank you for creating a wonderful working atmosphere that stimulated my creativity and thinking. I also thank Uwe John for his supervision and counsel throughout the whole time. Our inspiring discussions have always motivated me. Thanks for the academic freedom you two gave to me, and for the support I received from you with many respects, also aside from pure science. I would also like to thank professors Kai Bischof and Dieter Wolf-Gladrow for reviewing this thesis and being part of the promotion committee. Dieter, I have always enjoyed being part of the Biogeosciences group, which is not at last due to the unconventional way of your administration. I am indebted to this great group of people, for the open and collegial working environment and the fun that arose from it. Special thanks go to Albert Benthien, for his help and advice in so many things and the fun we had as fellow commuters. I would also like to thank Silke Thoms and Lena Holtz, who have not only sparked my interest for the dark arts of numerical modeling but also for the fruitful cooperation we had and will have. I want to express my gratitude to the technical staff of the group, especially Klaus-Uwe and Uli Richter, Jana Hölscher, Laura Wischnewski and all others that construct apparatuses, operate machines and run analyses. You all deserve great acknowledgments, for my work and that of others would not have been possible without you. I acknowledge the help and supervision I received from Scarlett Trimborn, Dedmer van de Waal and Sven Kranz, the senior (and partly former) members of the PhytoChange project. Thanks for your constructive criticism and the challenging discussions regarding methods, procedures and manuscript drafting. These were not always easy but have significantly influenced my interpretation and reasoning and improved the overall quality of my work. A lot of what I learned during the time of my PhD was due to the ‘never-let- loose’-attitude which I adapted from you, Sven. On the other side, I want to say ‘thank you’ to the younger members of the PhytoChange project, Clara Hoppe, Meri Eichner, Mirja Hoins, Doro Kottmeier and Tim Eberlein. It is a great pleasure and a delighting feeling to give to you what was previously given to me. Thank you Clara for being a kind and trustful office companion; our exciting discussions and arguments of whatever nature always gave me new insights or at least perspectives. Thank you Doro for being a dedicated and enthusiastic student, it is great to work with you and I look forward to our future explorations. I want to thank my cooperation partners, namely Colomban De Vargas and Ian Probert from the Biological Station Roscoff (France), for contributing their expertise to the 5 projects I conducted and the interesting discussion that arose. I also like to thank Peter von Dassow from the University of Santiago (Chile) for the great time we had creating loads of interesting data.
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