Isoprene Emission in Polytrichaceae Mosses
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Portland State University PDXScholar Dissertations and Theses Dissertations and Theses 5-24-2019 Isoprene Emission in Polytrichaceae Mosses Timea Deakova Portland State University Follow this and additional works at: https://pdxscholar.library.pdx.edu/open_access_etds Part of the Biology Commons Let us know how access to this document benefits ou.y Recommended Citation Deakova, Timea, "Isoprene Emission in Polytrichaceae Mosses" (2019). Dissertations and Theses. Paper 4984. https://doi.org/10.15760/etd.6860 This Dissertation is brought to you for free and open access. It has been accepted for inclusion in Dissertations and Theses by an authorized administrator of PDXScholar. Please contact us if we can make this document more accessible: [email protected]. Isoprene Emission in Polytrichaceae Mosses by Timea Deakova A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Biology Dissertation Committee: Sarah Eppley, Chair Todd Rosenstiel, Co-Chair Daniel Ballhorn Andrew Rice Erin Shortlidge Portland State University 2019 © 2019 Timea Deakova Abstract Our first aim was to identify and quantify Biological Volatile Organic Compound (BVOC) emissions, specifically emissions of isoprene, from the moss Polytrichum juniperinum during its earliest stage of life. Isoprene emission from mosses could be a significant component of the total global budget of BVOC emissions. Data concerning the spatial and temporal variability of these emissions are lacking due to poor characterization of the physical and biological factors controlling isoprene synthesis in both vascular and non-vascular plants. We found that P. juniperinum in its early life stage (protonema) can emit isoprene at detectable levels at day five after spore germination. These results suggest that protonema is capable of isoprene emission, shown here for the first time, and media affects emission rates. We saw a negative trend with leaf emergence and protonema growth. Chlorophyll fluorescence ratio had a positive correlation with isoprene emission, and isoprene emission was both temperature and light dependent in early protonema. Our second aim was to determine the effects of sex and sexual expression on isoprene emission in mosses. Sex is a biologically costly, but it can result in genetic adaptability and variety for resulting offspring. Using three Polytrichaceae species, we measured isoprene emission between plants sexually expressing and non-expressing and between male and female moss plants. We found that non-sexually expressing gametophytes had significantly higher isoprene emission than gametophytes expressing sex, suggesting that sex expression may be costly and that additional resources are i allocated to isoprene emission when plants are not reproducing. Males emitted higher levels of isoprene than females, but surprisingly this difference occurred only when plants were not expressing sex. We found species and sex-specific differences in chlorophyll fluorescence ratio (CFR) and relative electron transport rate (RETR). Our third aim was to investigate and categorize the effect of nitrogen addition on isoprene emission of P. juniperinum by creating an artificial nitrogen gradient with ammonium nitrate addition (NH4NO3). Current rates of anthropogenic N deposition are altering many biogeochemical processes. In these changing environments, increased nitrogen availability alters plant phenology, physiology, and the allocation of resources, but no information is available on whether additional N increases isoprene emissions in mosses. We used a manipulative experiment to measure the effects of nitrogen addition on moss isoprene emission, as well as on moss morphology and reproductive effort. We found site-specific differences in our morphological and physiological measurements. Isoprene emission was site, sex, and N addition specific, with the highest isoprene emission seen from our mid-level N addition, in both female- and male-majority pots. We found significant treatment and sex-specific interactions among our sites and within sites. We saw the highest reproductive effort counts from non-sexual expressive gametophytes, followed by high sporophyte count from female-majority pots. We also saw chlorophyll fluorescence ratio (CFR) differences between sites and treatments, but not between sexes. ii Acknowledgements I am extremely grateful for the advice, assistance, and encouragement I received from all my friends and colleagues during my graduate work. I was able to accomplish far more than I could have on my own. Undertaking this PhD has been a great life experience for me, and it would have been impossible to accomplish without the support and advice I received from many people. I would like first to thank my supervisor Dr. Sarah Eppley for her patience, mentorship for lending an ear to the many concerns and for the support, she provided me during this period. Her understanding and positive attitude was an important motivation for me to pursue my doctoral degree and I feel very fortunate to have had the opportunity to work with her support. I am also thankful to Dr. Rosenstiel; he introduced me to the wonderful world of mosses and gave me the opportunity to join his lab to explore them. I am grateful that he provided the insight I needed to explore isoprene emission in mosses. I also extend my thanks to my committee members, who have helped me reach the completion of this stage in my journey through academia with support and advice. Special thanks also to my friends and fellow lab mates and collaborators especially for their advice and friendship and support inside and outside the lab. They created a pleasant working environment that kept me motivated during my PhD while lending an ear to the inevitable gripes about the difficulties always experienced in the search for knowledge. I am especially thankful for Scott Kiel and his expertise in showing me the ropes in running the reduced gas analyzer. Thank you for sharing a passion of science with me. Thank you Scott! iii I would like to say heartfelt thanks to my family especially my children Annika and Mika, who have understood the long hours I have been working during their young lives. I am very thankful for my mother-in-law supported me taking care of my children when I was attending school. I would also like to thank my mother who has both supported me financially and emotionally through having children and going to school. I would like to thank my husband who was the first to show me how exciting science really is, how beautiful and elegant it can be, for sharing his sense of wonder at discovery a feeling that inspired hope, a hope that allowed me to summon my own strength to see this through. He was the first to believe in me, the first to recognize my abilities. I am thankful for his endless support in achieving my academic goals in his unflagging ability to convince me that all of this was possible even when I was not sure it was. Thank you, Doug, for your vision, energy, enthusiasm support in all the steps of my PhD. This work was funded by the National Science Foundation (NSF GRFP) Biological Science Grant 2015. Proposal Title: Moss Volatile Emissions: The role of BVOC in non-flowering sexual systems. I am extremely grateful to be awarded the NSF GRFP fund and without this award I would not have been able to complete my work. I want to thank additional funding from Sigma Xi, the American Association for the Advancement of Science Pacific Division. Lastly, I would also like to thank the Forbes- Lea research award and the Botanical Society Award that helped me get started on my initial data collection. iv Table of Contents Abstract ................................................................................................................................ i Acknowledgements ............................................................................................................ iii List of Tables ..................................................................................................................... vi List of Figures ................................................................................................................... vii Chapter 1 Introduction ..........................................................................................................................1 Chapter 2 Isoprene emission from Polytrichum juniperinum protonema. ...........................................9 Chapter 3 The role of isoprene in Polytrichaceae sexual systems .....................................................58 Chapter 4 The effect of nitrogen addition on moss morphology, sexual reproduction, and isoprene production. .................................................................................................127 Chapter 5 Conclusion .......................................................................................................................188 References ........................................................................................................................199 v List of Tables Table 2.1 Summary table of Polytrichum juniperinum populations used for protonema single spore and mixed spore experiments. .....................................................42 Table 3.1 Summary table of P. juniperinum populations used for non-sexually expressing isoprene emission (n=45). ..............................................................................100 Table 3.2 Summary table of P. juniperinum populations used for sexually