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Extracellular Vesicles and Nanoerythrosomes: the Hidden Pearls of Blood Products Academic Dissertations from The Finnish Red Cross Blood Service number 64 SAMI VALKONEN | Extracellular Vesicles and Nanoerythrosomes: The Hidden Pearls of Blood Products Blood Nanoerythrosomes: Pearls of Extracellular and Hidden The | Vesicles SAMI VALKONEN SAMI VALKONEN Extracellular Vesicles and Nanoerythrosomes: The Hidden Pearls of Blood Products ISBN 978-952-5457-48-3 (print) ISBN 978-952-5457-49-0 (pdf) ISSN 1236-0341 FRC BS: 64 http://ethesis.helsinki.fi Turku 2019 Painosalama Oy ACADEMIC DISSERTATION, NUMBER 64 Doctoral School of Health Sciences Doctoral Programme in Biomedicine Molecular and Integrative Biosciences Research Programme Faculty of Biological and Environmental Sciences University of Helsinki and Finnish Red Cross Blood Service EXTRACELLULAR VESICLES AND NANOERYTHROSOMES: THE HIDDEN PEARLS OF BLOOD PRODUCTS Sami Valkonen ACADEMIC DISSERTATION To be presented for public examination, with the permission of the Faculty of Biological and Environmental Sciences of the University of Helsinki, in Nevanlinna Auditorium of the Finnish Red Cross Blood Service, Kivihaantie 7, Helsinki, on 19th December 2019, at 12 noon. Helsinki 2019 ACADEMIC DISSERTATIONS FROM THE FINNISH RED CROSS BLOOD SERVICE, NUMBER 64 Supervisors Adjunct Professor Pia Siljander Faculty of Biological and Environmental Sciences University of Helsinki Doctor Saara Laitinen Department of Research & Development Finnish Red Cross Blood Service Thesis Committee Associate Professor Susanna Fagerholm Faculty of Biological and Environmental Sciences University of Helsinki Professor Kalle Saksela Faculty of Medicine University of Helsinki Pre-examiners Professor Andrew Devitt School of Life and Health Sciences Aston University Doctor Paul Harrison College of Medical and Dental Sciences University of Birmingham Opponent Associate Professor Kenneth Witwer Department of Molecular and Comparative Pathobiology; Department of Neurology Johns Hopkins University School of Medicine Custos Professor Kari Keinänen Faculty of Biological and Environmental Sciences University of Helsinki The Faculty of Biological and Environmental Sciences uses the Urkund system (plagiarism recognition) to examine all doctoral dissertations. ISBN 978-952-5457-48-3 (print) ISBN 978-952-5457-49-0 (pdf) ISSN 1236-0341 http://ethesis.helsinki.fi Turku 2019 Painosalama Oy “All the adversity I've had in my life, all my troubles and obstacles, have strengthened me... You may not realize it when it happens, but a kick in the teeth may be the best thing in the world for you.” -Walt Disney To the closest ones to my heart TABLE OF CONTENTS TABLE OF CONTENTS ..............................................................................................i ORIGINAL PUBLICATIONS ................................................................................... ii PERSONAL CONTRIBUTION ............................................................................... iii PUBLICATIONS NOT INCLUDED IN THE THESIS ........................................ iv ABBREVIATIONS ....................................................................................................... v ABSTRACT ................................................................................................................. vi 1 BACKGROUND ................................................................................................. 1 1.1 From donated blood to transfusable products .................................. 1 1.1.1 Red blood cells, dedicated servants of gas exchange............................ 3 1.1.2 Platelets, more than haemostasis ......................................................... 4 1.2 Extracellular vesicles ............................................................................... 8 1.2.1 The significance of red blood cell and platelet-derived extracellular vesicles .............................................................................. 9 1.2.2 Cargo-based signalling in extracellular vesicles ................................ 11 1.2.3 Extracellular vesicle assessment ........................................................ 14 1.2.3.1 Isolation of extracellular vesicles ............................................... 16 1.2.3.2 Quantification and characterisation of extracellular vesicles .. 18 1.2.4 Beyond biology, toward applications and commercial value ............. 22 1.3 Membrane lipid signalling in platelets .............................................. 25 2 AIMS OF THE STUDY ................................................................................... 29 3 MATERIALS AND METHODS ..................................................................... 30 3.1 Blood products ......................................................................................... 30 3.2 Extracellular vesicle isolation ............................................................. 31 3.3 Reference material production ........................................................... 31 3.4 Reference material characterisation ................................................. 31 3.5 Platelet activation assessment ............................................................ 32 3.6 Membrane lipid and fatty acid analysis ............................................ 32 3.7 Lipid mediator and pathway marker analysis ................................ 32 3.8 Flow cytometry ........................................................................................ 33 3.9 Nanoparticle tracking analysis ........................................................... 33 3.10 Western blot.............................................................................................. 33 3.11 Statistical analysis .................................................................................. 34 4 RESULTS .......................................................................................................... 35 4.1 Nanoerythrosomes as a reference material ..................................... 35 4.2 Extracellular vesicles and platelet activation ................................. 38 4.3 Lipid-mediated signalling of extracellular vesicles ....................... 40 5 DISCUSSION ................................................................................................... 45 5.1 On the road to traceable extracellular vesicle assessment .......... 45 5.2 The extracellular vesicles of platelet concentrates: The good, the bad, or the ugly? ............................................................ 52 5.3 Future perspectives ................................................................................ 60 6 FINAL CONCLUSIONS ................................................................................. 62 7 ACKNOWLEDGEMENTS .............................................................................. 64 8 REFERENCES ................................................................................................. 68 9 APPENDICES .................................................................................................. 95 i ORIGINAL PUBLICATIONS This thesis is based on the following publications that are referred to in the text by their roman numerals (Studies I-III). In addition, some unpublished results are presented. I Valkonen S, van der Pol E, Böing A, Yuana Y, Yliperttula M, Nieuwland R, Laitinen S, Siljander PRM. Biological reference materials for extracellular vesicle studies. Eur J Pharm Sci 2017;98:4–16. II Valkonen S, Mallas B, Impola U, Valkeajärvi A, Eronen J, Javela K, Siljander PRM, Laitinen S. Assessment of Time-Dependent Platelet Activation Using Extracellular Vesicles, CD62P Exposure, and Soluble Glycoprotein V Content of Platelet Concentrates with Two Different Platelet Additive Solutions. Transfus Med Hemotherapy 2019;46:267–75. III Valkonen S§, Holopainen M§, Colas RA, Impola U, Dalli J, Käkelä R, Siljander PRM, Laitinen S. Lipid mediators in platelet concentrate and extracellular vesicles: Molecular mechanisms from membrane glycerophospholipids to bioactive molecules. Biochim Biophys Acta - Mol Cell Biol Lipids 2019;1864:1168–82. § = equal contribution The publications have been included in the thesis as appendix with kind permission from Elsevier (Amsterdam, Netherlands (Studies I, III)) and S. Karger AG (Basel, Switzerland (study II)). ii PERSONAL CONTRIBUTION I The author participated in the design of the study, conducted the literature review, developed and sent the questionnaire, recorded and analysed the questionnaire results, optimised the reference material production and conducted the majority of the characterisation (size and particle concentration determination, stability, flow cytometry, Nanoparticle tracking analyser (NTA) comparison), data analysis, and result interpretation. The final manuscript was written mostly by the author and it was critically revised by the co-authors. II The author participated in the study design, performed the NTA data collection and analysis, performed the statistical analysis throughout the results and interpreted the results together with supervisors. The final manuscript was written mostly by the author, who also participated in the manuscript revision. III The author participated in the study design, performed the NTA, phospholipid, and Western blot data collection as well as data analysis. The author interpreted the results with the co-authors, mostly wrote the final manuscript and participated in the manuscript revision. iii PUBLICATIONS NOT INCLUDED IN THE THESIS In addition to the publications included in the thesis, the author has contributed to the
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