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Genome Size and Phenology 14 Stomata 16 Stomata: the Role of Genome Size Bibliographic identification Author: Mgr. Pavel Veselý Title of the thesis: Have the largest plant genomes evolved as an adaptati- on to geophytic strategy? Study programme: Biology Study field: Botany Supervisor: doc. RNDr. Petr Bureš, Ph.D. Academic year: 2014–2015 Keywords: Genome size evolution, AT/GC genomic ratio, geophytes, life-form, storage organ, tuber, bulb, rhizome, phenolo- gy, ephemeroids, ecology, stomatal size, stomatal speed, seed mass i Bibliografická identifikace Autor: Mgr. Pavel Veselý Název práce: Vznikly největší genomy rostlin jako součást adaptace na geofytní strategii? Studijní program: Biologie Studijní obor: Botanika Školitel: doc. RNDr. Petr Bureš, Ph.D. Akademický rok: 2014–2015 Klíčová slova: Evoluce velikosti genomu, AT/GC genomový poměr, geo- fyty, životní forma, zásobní orgán, hlíza, cibule, odde- nek, fenologie, efemeroidy, ekologie, velikost průduchů, rychlost průduchů, hmotnost semen ii Abstract For several decades, the genome size and its selective significance is a subject of re- search, nevertheless, some questions still remain unanswered. The genome size af- fects a number of plant traits (e. g. stomatal size, maximum height, seed mass) and has a series of physiological (e. g. water use effiiny ad eooia (.g phenology minimum generation time, life forms) implications. However, it is not clear how do these relationship function for species with giant genomes and if and how are these species limited. There is also a question what consequences a different GC base content in genomes can have and if the base ratio relates somehow to genome size. Thus I focused my dissertation thesis on geophytes which due to frequent occurrence of large genomes appear to be a suitable model group for testing the mentioned relationships. Acquired data show that evolution of large genomes in geophytes is closely as- sociated with earlier flowering, larger stomata and shorter growing season. The key to such constraint could be seen in a physiology of stomata. Based on the measurements of more than thirty pairs of related species with contrasting ge- nome and stomatal sizes, larger stomata showed to be slower than smaller ones which resulted in higher transpirative water loss. The increase of genome size was observed not only in spring geophytes. Comparison of 47 pairs of geophytes and their non-geophytic relatives confirms that the transition to geophytic strat- egy leads to an evolution of larger genomes that can be accounted to the energy reserves in the subterranean organ. The genomic GC base composition of geo- phytes does not tell much about their ecology, still it has a unimodal relationship with genome size. There could exist a general constraint on organization of DNA which would prevent a higher GC contents in large genomes. The seed mass of geophytes did not show significant link to genome size, yet it relates to ecology which perhaps plays a more important role in selection and masks the potential effect of the genome size. iii Anotace Velikost genomu u rostlin a její selektivní význam je již několik desetiletí předmě- tem výzkumu, přesto však některé otázky stále zůstávají nezodpovězeny. Velikost genomu ovlivňuje množství znaků na rostlinách (např. velikost průduchů, maxi- mální výška, hmotnost semen) a má celou řadu fyziologických (např. hospodaření s vodou) a ekologických (např. fenologie, minimální generační doba, životní formy) důsledků. Není však jasné, jak tyto vztahy fungují u druhů s obřími genomy a zda a případně jak jsou tyto druhy limitovány. Otázkou je též, jaký vliv může mít rozdíl- né zastoupení GC bází v genomech a zda tento poměr bází má nějakou souvislost s velikostí genomu. Proto jsem se ve své dizertační práci zaměřil na geofyty, které se díky častému výskytu velkých genomů jeví jako vhodná modelová skupina k tes- tování zmíněných vztahů. Získaná data ukazují, že evoluce velkých genomů u geofytů je těsně spojena s brzkým kvetením, většími průduchy a kratší vegetační sezónou. Klíčem k pocho- pení takového omezení by mohla být fyziologie průduchů, kdy na základě měření více než třiceti párů příbuzných druhů s rozdílnou velikostí genomu a průduchů se velké průduchy ukázaly být proti menším pomalejší, což znamenalo větší ztrátu vody transpirací. Nárůst velikosti genomu nebyl pozorován jen u jarních geofytů. Srovnání 47 párů geofytů a příbuzných negeofytů potvrzuje, že přechod na geo- fytní strategii vede k evoluci větších genomů, což lze připisovat zásobám energie v podzemních orgánech. Obsah GC bází u geofytů sice příliš nevypovídá o jejich ekologii, ale má unimodální vztah k velikosti genomu. Mohla by tedy existovat obecnější příčina v uspořádání DNA ve velkých genomech, která by vyšší podíl GC bází vylučovala. Hmotnost semen geofytů neukázala významný vztah k velikosti genomu, zato však souvisí s ekologií, která patrně hraje významnější roli v selekci a zakrývá možný vliv velikosti genomu. iv © Pavel Veselý, Masarykova univerzita, 2015 v Contents – Obsah Acknowledgements 4 Preface 5 Genome size 7 What is a genome size? ............................................................................................................ 7 Why to study genome size? .................................................................................................. 7 When the size matters more than the content ................................................ 8 Genome size and life forms 9 Genome sizes in particular life forms ....................................................................... 9 Geophytism and genome size ..........................................................................................10 Geophytism as a strategy ....................................................................................................12 Genome size and phenology 14 Stomata 16 Stomata: the role of genome size ...............................................................................16 Stomata, their size and regulation ............................................................................18 Genome size and DNA base composition 25 Genome size and seed mass 28 Summary of used methods 30 Genome size and GC content ..........................................................................................30 Chromosomal counts and ploidy levels ................................................................31 Phenology of flowering and duration of growing season ..................31 Stomatal size .....................................................................................................................................32 Stomatal closure speed .........................................................................................................32 Seed mass ............................................................................................................................................34 Ecological characteristics ...................................................................................................34 Statistics ................................................................................................................................................34 Conclusions 36 Poděkování 38 Místo úvodu 39 Velikost genomu 41 Co je to velikost genomu? ...................................................................................................41 Proč se zajímat o velikost genomu? .........................................................................41 1 Když více záleží na množství než na obsahu ...................................................42 Velikost genomu a životní formy 43 Vztah velikosti genomu a životních forem ........................................................43 Geofytismus a velikost genomu ....................................................................................44 Geofyt jako strategie ................................................................................................................46 Velikost genomu a fenologie geofytů 48 Průduchy 50 Průduchy: role velikosti genomu .................................................................................50 Velikost průduchů a jejich regulace .........................................................................52 Obsah GC bází v genomu 58 Velikost genomu a hmotnost semen 61 Souhrn použitých metod 63 Velikost genomu a obsah GC bází ..............................................................................63 Chromozómové počty a stupeň ploidie .................................................................64 Fenologie kvetení a délka vegetační sezóny ...................................................64 Velikost průduchů ........................................................................................................................65 Rychlost zavírání průduchů ..............................................................................................65 Hmotnost semen ...........................................................................................................................67 Ekologické charakteristiky ................................................................................................67 Statistika ...............................................................................................................................................67 Shrnutí 68 References – Literatura 69 Curriculum vitae 76 Životopis 80 Author contribution to presented papers 83 Paper 1 84 Paper 2 102 Paper 3 116 2 Masaryk University Faculty of Science Department of Botany and Zoology Have the Largest Plant Genomes Evolved as an Adaptation
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