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Reproduction and Hybridization in Ferns

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Reproduction and Hybridization in Ferns School of Doctoral Studies in Biological Sciences University of South Bohemia in České Budějovice Faculty of Science Reproduction and hybridization in ferns Ph.D. Thesis Mgr. Ondřej Hornych Supervisor: RNDr. Libor Ekrt, Ph.D. České Budějovice 2020 This thesis should be cited as: Hornych, O., 2020: Reproduction and hybridization in ferns. Ph.D. Thesis Series 2020, No. 22. University of South Bohemia, Faculty of Science, School of Doctoral Studies in Biological Sciences, České Budějovice, Czech Republic, 162 pp. Annotation This thesis studies several aspects of fern reproduction and hybridization. Aspects of multiple stages in the fern life cycle are analyzed. Specifically, rates of spore abortion and antheridiogen usage are assessed and compared between various groups. Own cultivation and spore assessment data are compared extensively with existing literature. Finally, the rate of hybridization in natural populations and the symmetry of hybridization is analyzed in buckler ferns (Dryopteris carthusiana group). Declaration [in Czech] Prohlašuji, že svoji disertační práci jsem vypracoval samostatně pouze s použitím pramenů a literatury uvedených v seznamu citované literatury. Prohlašuji, že v souladu s § 47b zákona č. 111/1998 Sb. v platném znění souhlasím se zveřejněním své disertační práce, a to v úpravě vzniklé vypuštěním vyznačených částí archivovaných Přírodovědeckou fakultou elektronickou cestou ve veřejně přístupné části databáze STAG provozované Jihočeskou univerzitou v Českých Budějovicích na jejích internetových stránkách, a to se zachováním mého autorského práva k odevzdanému textu této kvalifikační práce. Souhlasím dále s tím, aby toutéž elektronickou cestou byly v souladu s uvedeným ustanovením zákona č. 111/1998 Sb. Zveřejněny posudky školitele a oponentů práce i záznam o průběhu a výsledku obhajoby kvalifikační práce. Rovněž souhlasím s porovnáním textu mé kvalifikační práce s databází kvalifikačních prací Theses.cz provozovanou Národním registrem vysokoškolských kvalifikačních prací a systémem na odhalování plagiátů. České Budějovice, 1.12.2020 .............................................. Ondřej Hornych Financial support This thesis was supported by the Czech Science Foundation (grant number 19-17379S) and by the Grant Agency of the University of South Bohemia (grant number 022/2019/P). Acknowledgements I would like to thank my supervisor Libor Ekrt for his continuous support with my research endeavors. He spent considerable time helping me with all the aspects of my research, teaching me methods and plant identification, spending time in the field with me, providing me with his various fern specimens, obtaining financial support and helping me with all aspects paper preparation. All throughout he also always approached me as a friend. I am also very grateful to my colleagues. To Weston Testo and Emily Sessa for their immense help, support and inspiration during my stay at the University of Florida and during the preparation of the antheridiogen manuscript. To Jiří Košnar for supporting me throughout the not always successful experiments with fern cultivations and guiding my molecular laboratory efforts. To Petr Koutecký for his invaluable advice in statistical analyses, flow cytometry, experimental design and paper writing. To my coauthors and department colleagues in general, for their help in many aspects, in science, teaching, learning and navigating administrative affairs. Finally, my thanks go to my close friends and family, for their support in the past four years and more. List of papers and author’s contribution The thesis is based on the following papers (listed chronologically): Hornych O, Ekrt L. 2017. Spore abortion index (SAI) as a perspective tool of evaluation of spore fitness in ferns: An insight into sexual and apomictic species. Plant Systematics and Evolution 303: 497–507. (IF = 1.15) OH and LE conceived the study, collected samples and contributed to the writing of the manuscript, OH performed SAI assessment and statistical analyses. Hornych O, Ekrt L, Riedel F, Koutecký P, Košnar J. 2019. Asymmetric hybridization in Central European populations of the Dryopteris carthusiana group. American Journal of Botany 106: 1477–1486. (IF = 2.84) OH and LE conceived the study. OH, FR, and LE collected field data. All authors performed laboratory analyses, contributed to writing the manuscript, and gave final approval for publication. Hornych O, Testo WL, Sessa EB, Watkins JE Jr, Campany CE, Pittermann J, Ekrt L. 2020. Insights into the evolutionary history and widespread occurrence of antheridiogen systems in ferns. New Phytologist In press, doi: 10.1111/nph.16836. (IF = 8.51) OH, WLT, JEW and LE designed the study; OH, WLT, CEC and JP conducted the cultivation experiments; OH compiled the meta-analysis list; and OH, WLT and EBS analyzed the data. All authors contributed to the writing of the manuscript. Contents Chapter 1: General introduction ........................................................................................ 1 Introducing the life of plants in general, and ferns in particular. .................................. 1 The life cycle of land plants ...................................................................................... 1 The specifics of ferns ................................................................................................ 1 The spore, the means of fern travel and the check of genetic integrity ......................... 3 Spores as means of dispersal ..................................................................................... 3 Spore size, shape and color ....................................................................................... 3 Spore abortion ........................................................................................................... 4 Summary ................................................................................................................... 5 The gametophyte, the short and turbulent stage in a fern's life ..................................... 6 Factors affecting gametophyte growth ...................................................................... 6 Gametophyte ontogeny ............................................................................................. 6 Antheridiogens .......................................................................................................... 7 Choosing a mate ........................................................................................................ 8 Apomixis ................................................................................................................... 9 Summary ................................................................................................................... 9 The sporophyte, the conspicuous and resilient stage in a fern's life ............................ 10 The legacy of sporophytes ...................................................................................... 10 Sporogenesis ........................................................................................................... 11 Apomictic hybrids ................................................................................................... 12 Summary ................................................................................................................. 12 Aims of the thesis ........................................................................................................ 13 References ................................................................................................................... 14 Chapter 2: Spore abortion index (SAI) as a perspective tool of evaluation of spore fitness in ferns: An insight into sexual and apomictic species.................................................... 21 Abstract ....................................................................................................................... 22 Introduction ................................................................................................................. 22 Materials and methods ................................................................................................ 24 Plants used in the study ........................................................................................... 24 Evaluation of spore abortion rates ........................................................................... 24 Data analyses........................................................................................................... 26 Results ......................................................................................................................... 28 Determining aborted spores .................................................................................... 28 Spore abortion index ............................................................................................... 28 Comparing SAI of different groups ........................................................................ 30 Optimization of SAI assessment method ................................................................ 30 Discussion ................................................................................................................... 34 Effects of reproduction mode on spore abortion ..................................................... 34 Effects of ploidy levels on spore abortion ............................................................... 35 Different rates of spore abortion among genera .....................................................
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