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Juncus Effusus ) Relevant for Natural Degradation of Contaminants

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Juncus Effusus ) Relevant for Natural Degradation of Contaminants Evolutionary potential in functional traits of a wetland macrophyte ( Juncus effusus ) relevant for natural degradation of contaminants Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften (Dr. rer. nat.) der Naturwissenschaftlichen Fakultät I – Biowissenschaften – der Martin-Luther-Universität Halle-Wittenberg, vorgelegt von Frau Jennifer Born (MSc Biol.) geboren am 18.09.1989 in Leisnig Referees Dr. Stefan G. Michalski (Helmholtz Centre for Environmental Research – UFZ, Department Community Ecology) Prof. Dr. Helge Bruelheide (Martin-Luther-University Halle-Wittenberg, Institute of Biology / Geobotany and Botanical Garden) Prof. Dr. Frank H. Hellwig (Friedrich-Schiller-University Jena, Institute of Systematic Botany) Thesis defence 22.01.2019 TABLE OF CONTENTS SUMMARY ............................................................................................................................ 4 ZUSAMMENFASSUNG ........................................................................................................ 6 CHAPTER 1 General Introduction Wetlands and their ecological relevance ...........................................................................11 The nitrogen cycle: processes, agents and human influence ............................................12 Plant functional traits and trait variability ...........................................................................13 Evolutionary and ecological genetics ................................................................................15 Quantitative genetic diversity in functional traits ................................................................17 DNA-based marker systems to assess genetic diversity ...................................................19 Evolutionary potential, genetic diversity and local adaptation ............................................20 Juncus effusus – a representative wetland plant ...............................................................22 Objectives and thesis outline ............................................................................................24 CHAPTER 2 Strong Divergence in Quantitative Traits and Plastic Behavior in Response to Nitrogen Availability among Provenances of a Common Wetland Plant ..............................................25 CHAPTER 3 Trait Expression and Signatures of Adaptation in Response to Nitrogen Addition in the Common Wetland Plant Juncus effusus ...............................................................................46 CHAPTER 4 Genetic Analyses Reveal That Multi-Facetted Processes are Shaping the Intraspecific Differentiation Patterns of the Widespread Wetland Plant Juncus effusus in Europe ............70 CHAPTER 5 Synthesis .......................................................................................................92 Assessing the importance of intraspecific variability ..........................................................94 Evidence of selection ........................................................................................................98 Implications for applied approaches ................................................................................ 100 Lineage divergence, isolation and genetic drift in Juncus effusus ................................... 102 Coexistence of lineages .................................................................................................. 104 Concluding remarks ........................................................................................................ 106 SUPPORTING INFORMATION .......................................................................................... 108 REFERENCES ................................................................................................................... 111 DANKSAGUNG ................................................................................................................. 125 APPENDIX ......................................................................................................................... 127 SUMMARY SUMMARY Nutrients such as nitrogen and phosphorus are natural components of aquatic ecosystems and essential for plant growth. However, if progressive intensification of agriculture and urbanization cause too many nutrients to enter the ecosystem, then air and water can be polluted. Wetland ecosystems provide the important ecosystem service of natural degradation and retention of such excessive substances. The remediation potential of wetland plants and associated microorganisms has increased interest in the rehabilitation of lost and degraded wetlands. In addition, wetland constructions have frequently been used as cost-effective and sustainable biotechnologies to compensate for the loss of wetlands in the last century. Plant traits can have a direct or indirect impact on wetland ecosystems through accumulation, metabolism and microbial transformation processes in the soil. The influence of plants’ functional traits and their interaction with the soil microbiome in the reduction of wetland pollutants have been well investigated with respect to the among-species variability. However, the extent of the intraspecific variability of these functional traits may be as important as interspecific variability in relation to ecosystem functions. Understanding the causes and consequences of intraspecific genetic variation related to nutrient degradation processes is fundamental for predicting evolutionary responses, but it can also aid applied approaches in wetland restoration and construction. Applying a combination of quantitative and molecular genetics, the aim of the thesis is to quantify intraspecific variability in functional traits that are potentially relevant for the degradation of nitrogen compounds, and to investigate its genetic basis and response to selection. In our research, we used the wetland plant Juncus effusus , which has been established as a model plant in basic and applied research on wetland ecosystems. To understand whether and how quantitative genetic variation is expressed depending on nitrogen availability and how it is structured within and among natural European populations, we used common garden studies to evaluate and compare trait variation across multiple provenances, lineages and treatments. Microsatellites were used to study genetic diversity and population - 4 - SUMMARY structure to investigate the different environmental forces and their relative contributions to adaptation. Then we investigated possible relationships between genetic diversity and environmental factors such as climate and soil conditions. Furthermore, we asked whether trait expression shows signatures of local adaptation by comparing quantitative genetic trait divergence and neutral molecular markers (QST – FST comparisons) and relating trait variation to soil conditions of the plant’s origin. We quantified a strong plastic behavior in response to nitrogen availability in all measured traits. The genetic variation of functional mean traits was strongly pronounced among populations, but very low at population level. We could not find a significant link between nitrogen availability and genetic trait expression. Differentiation of quantitative traits was substantial but did not exceed neutral expectations, as detected in QST – FST comparisons. However, adaptive trait divergence in response to environmental soil conditions might be present, which is indicated by significant trait clines. Molecular analyses showed moderate genetic diversity and very strong genetic differentiation among populations. These findings confirmed our hypothesis, which is based on the life history of species Juncus effusus , as a successful colonizer with a predominantly selfing mating system. A Bayesian analysis of population structure across the European range showed that J. effusus comprises three genetically well-differentiated lineages that partly separate longitudinally but also co-occur sympatrically, probably as a result of glacial lineage diversification and secondary contact. The lineage Eff3 differed substantially from the other lineages by clines in population genetic diversity with environmental conditions and by a dominant occurrence at higher altitudes suggesting more particular ecological responses compared to lineages Eff1 and Eff2. In summary, however, we found only little and inconsistent evidence of selective mechanisms varying between the lineages, which supports the idea that lineages within J. effusus are the primary results of neutral divergence. - 5 - ZUSAMMENFASSUNG ZUSAMMENFASSUNG Nährstoffe wie Stickstoff und Phosphor sind natürliche Bestandteile aquatischer Ökosysteme und für das Pflanzenwachstum unerlässlich. Wenn jedoch zu viele Nährstoffe in das Ökosystem gelangen, können Luft und Wasser verschmutzt werden. Diese Verschmutzung wird hauptsächlich durch die fortschreitende Intensivierung der Landwirtschaft und Urbanisierung verursacht. Feuchtgebietsökosysteme leisten einen wichtigen Beitrag durch natürlichen Abbau und Rückhaltung solch überschüssiger Nährstoffe. Das Sanierungspotential von Feuchtgebietspflanzen und den im Boden lebenden Mikroorganismen hat das Interesse an der Sanierung von verloren gegangenen und degradierten Feuchtgebieten erhöht. Darüber hinaus werden künstlich angelegte Feuchtgebietsanlagen häufig als kostengünstige und nachhaltige Biotechnologie eingesetzt, um den Verlust von natürlichen Feuchtgebieten auszugleichen. Pflanzeneigenschaften können die Ökosystemprozesse von Feuchtgebieten
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