phosphomimicking nor kinase-inactivation assay in HEK cells were performed with this residue.
However, in an in vitro assay, T174 was highly phosphorylated by membrane kinases, and mildly by soluble kinases. The peptide containing this serine was also phosphorylated in in vitro peptide kinase assays. Phosphomimicking suggested that phosphorylation of T174 greatly activates PaRBOH1 enzyme activity. As MF membranes from developing xylem were used as a source of kinases, this suggests that such phosphorylation can occur during normal xylem development. These results suggest that kinases in the MF membranes could be interesting candidates for PaRBOH1 activation. There were 50 kinases detected altogether in the proteomic analysis (II), of which 46 were detected in Norway spruce developing xylem samples.
Among these kinases, a probable receptor-like protein kinase (MA_118589g0010) was co-expressed with monolignol biosynthesis genes in all the datasets (II). It is tempting to speculate that this kinase may be responsible for regulating enzymes involved in lignification. However, any discussion about a kinase for T174 without any evidence is extremely hypothetical. In the future, further experiments would be required to identify a kinase for PaRBOH1 activity regulation.
studies with some of the membrane proteins detected here, a study on membrane lipidomes and investigations on interactions of the membrane components will all hopefully be forthcoming as part of lignin biosynthesis research conducted in the future.
ACKNOWLEDGEMENTS
This work was carried out in the Organismal and Evolutionary Biology Research Programme of University of Helsinki. In addition, majority of the laboratory work was physically conducted in the Department of Agriculture and Forestry. I want to thank both laboratories and University of Helsinki for enabling my studies and research. For the funding I would like to thank Viikki Graduate School in Molecular Biosciences (VGSB) and later Doctoral Programme in Integrative Life Science (ILS), as well as Jenny and Antti Wihuri foundation.
I want to express my gratitude to my supervisors Docent Anna Happonen and Professor Kurt Fagerstedt for their time, support, and forbearance. Anna, you have taught me so much of research in practice and patiently pushed me forward. Your passion for science has been contagious and given me energy when I have felt that my project is overwhelmingly difficult. Kurt, you have always met me with such kindness, and when I think that something is impossible, you find another perspective. And thank you with your help with all the bureaucracy during different steps of this project.
I would like to thank Docent Olga Blokhina for being such a great colleague and patiently listening to me when I preached about multiple conclusions that I made of my results. You always meet other people with kindness and respect, and despite being such an experienced scientist yourself, you have encouraged me by treating me like an expert, too.
I would like to thank Professor Teemu Teeri, for the tough questions in the “phenolics meetings” and great ideas with various experimental setups. Thank you also for Professor Juha Voipio for giving membrane- and cell physiology perspective for my results. Thank you for Associate professor Saijaliisa Kangasjärvi for the help in starting the pre-examination process.
Thank you also for Professor Lacey Samuels for agreeing to be my opponent, and preliminary examiners Associate professor Totte Niittylä and Dr. Esther Novo-Uzal for their comments on my thesis.
I would also like to thank all the collaborators of the projects involved in all the three projects within this thesis. Also, thank you for the friends with whom I have had a great privilege to work with: Teresa Laitinen, Tanja Paasela and Kaija Porkka. You have all helped a lot and been important support! Yaseen Mottiar, thank for your help with the final steps of my thesis! Thank you Marja Huovila, Anu Rokkanen, Marjo Kilpinen, and especially Eija Takala, who has helped me in various steps in my experiments. In addition, thank you for wonderful former members of our group, Kaloian Nickolov and Adrien Gauthier as well as Sharif, Eliisa, Luis, and Amanda, you have all been so wonderful to work with. Special thanks for Kaj-Roger Hurme from the isotope lab for such a nice working atmosphere and various stories and encouragement. Thank you Soile Jokipii-Lukkari for the discussion about spruce monolignol biosynthesis genes.
Million times thank you for the research-friends/officemates for scientific and non- scientific discussions for Marcelina, Eeva, Sanna, Tarja, Kean-Jin, Hany, Milla, Nashmin, Marja, Jasmin, Nerea, Santeri, and many more.
Of the very non-scientific stuff, that has kept me sane during these years I want to thank all my dear friends! A big heart-shaped thank you for all the branches of my family as they have tolerated all my PhD student symptoms for such a long time! Thank you for the whole nearly-in-law family for all the fun times. Thank you for my mom and dad for never questioning my career decisions, for support, friendship, and for the help with kids! And thank you for my sister, Tuuli, for being my friend and always being on my side. And finally thank you Ville for putting my PhD into perspective with your genius suggestions to my problems, and the various things that have happened in our lives during these years. Thank you Antto and Lyyti for all the happiness and love you have given me!
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