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Chapter 2 Effects of Leaf Water Evaporative Research Collection Doctoral Thesis Partial transfer of leaf water ²H-enrichment and variable biosynthetic fractionation affect the leaf wax n-alkane δ²H values in grasses Author(s): Gamarra, Bruno Publication Date: 2016 Permanent Link: https://doi.org/10.3929/ethz-a-010713453 Rights / License: In Copyright - Non-Commercial Use Permitted This page was generated automatically upon download from the ETH Zurich Research Collection. For more information please consult the Terms of use. ETH Library DISS. ETH No. 23286 Partial transfer of leaf water 2H-enrichment and variable biosynthetic fractionation affect the leaf wax n-alkane δ 2H values in grasses A thesis submitted to attain the degree of DOCTOR OF SCIENCES of ETH ZURICH (Dr. sc. ETH Zurich) presented by BRUNO GAMARRA MSc ETH Environ. Sc., ETH Zurich born on 23.06.1982 of Peruvian nationality accepted on the recommendation of Prof. Dr. Ansgar Kahmen Prof. Dr. Tim Eglinton Dr. Rolf Siegwolf 2016 Table of contents i Table of contents Summary ................................................................................................................................... 1 Zusammenfassung .................................................................................................................... 5 Chapter 1 ................................................................................................................................... 9 General introduction Chapter 2 ................................................................................................................................. 25 Effects of leaf water evaporative 2H-enrichment and biosynthetic fractionation on leaf wax n-alkane δ2H values in C3 and C4 grasses Chapter 3 ................................................................................................................................. 67 Low secondary leaf wax n-alkane synthesis on fully mature leaves of C3 grasses grown at low and high humidity Chapter 4 ................................................................................................................................. 95 Concentrations and δ2H values of cuticular n-alkanes vary significantly among plant organs, species and habitats in grasses from an alpine and a temperate European grasslands Chapter 5 ............................................................................................................................... 129 Conclusions Acknowledgements ............................................................................................................... 137 ii Table of contents Summary 1 Summary Leaf wax n-alkanes are long chained hydrocarbons contained in the cuticle of terrestrial plants. Leaf wax n-alkane δ2H values have been successfully used to study the hydrological cycle, plant-water and plant-carbon relationships. However, the physiological and biochemical drivers that shape leaf wax n-alkane δ2H values are not completely understood. It is particularly unclear, why n-alkanes in grasses are typically 2H-depleted compared to dicotyledonous plants and why C3 grasses are 2H-depleted compared to C4 grasses. Also, the timing of n-alkane synthesis and the de novo synthesis of n-alkanes in fully matured leaves are still matter of debate. On top, calibration studies designed to resolve sources of heterogeneity in n-alkane δ2H values have exclusively focused on n-alkanes derived from leaves. To solve these uncertainties three studies aimed to explore how leaf wax n-alkane δ2H values in grasses are influenced by plant physiological and biochemical processes. In the first study the effect of leaf water evaporative 2H-enrichment (LW Δ2H) on n-alkane δ2H values was quantified for a range of C3 and C4 grasses that were grown in climate chambers under controlled environmental conditions. It was found that only a fraction of the LW Δ2H is imprinted on the n-alkane δ2H values in C3 and C4 grasses. It was also detected that the biosynthetic hydrogen isotope fractionation (εbio) was different for C3 and C4 grasses. As such, differences between leaf wax n-alkane δ2H values of C3 and C4 grasses would not be 2 driven by LW Δ H but largely the result of systematic differences in εbio between these two plant groups. 2 Summary In the second study the timing of the leaf wax n-alkane synthesis was investigated. Specifically the secondary leaf wax n-alkane synthesis was tested in mature leaf blades of C3 grass species. The experiments showed an incorporation of hydrogen from highly 2H- enriched irrigation water into the wax n-alkanes from mature leaves. Mature grass leaves continued the synthesis of wax n-alkanes after leaf emergence. The rate of secondary n- alkanes synthesis was, however, relatively low and varied among species from 0.09% to 1.09% per day. As such, leaf wax n-alkane δ2H values would be determined mainly by environmental and physiological conditions in the beginning of the life of a leaf. In the third study n-alkane concentration and δ2H values of different grass organs were surveyed. Leaves, sheaths, stems, inflorescences and roots were sampled from a total of 15 species of C3 grasses in temperate and alpine grasslands in Switzerland. It was detected that inflorescences had typically much larger n-alkane concentrations compared to other organs while roots had very low n-alkane concentrations. The δ2H values of the carbon autonomous plant organs such as leaves, sheaths and stems were in general more negative compared to the non-carbon autonomous organs such as inflorescences and roots. Variable n-alkane δ2H values in different plant organs could be the result of differences in the H- NADPH biosynthetic origin in response to the carbon autonomy of the plant organ. Overall, this thesis brings new insights into the natural variability of leaf wax n-alkane δ2H values in grasses. The incomplete transfer of LW Δ2H to the leaf wax n-alkanes δ2H values in grasses can explain why grasses are typically 2H-depleted compared to dicotyledonous plants. The low secondary leaf wax n-alkane synthesis in grass leaves after maturity suggests that in general leaf wax lipid δ2H values do not record environmental and plant physiological processes beyond leaf maturity. Finally, a different εbio between C3 and C4 grasses and also between grass organs suggests that n-alkane δ2H values have a great Summary 3 potential as indicator of changes in plant carbon autonomy. This has important implications for the interpretation of n-alkane δ2H values in plant physiology and paleoecology. 4 Summary Zusammenfassung 5 Zusammenfassung Blattwachs n-Alkane sind langkettige Kohlenwasserstoffe die in der Epidermis von Landpflanzen vorkommen. δ2H-Werte von Blattwachs n-Alkanen wurden bisher erfolgreich verwendet um den Wasserkreislauf und die Beziehungen zwischen der Pflanze und dem Wasser sowie dem Kohlenstoff zu untersuchen. Dennoch sind die Auswirkungen von physiologischen und biochemischen Prozessen auf die δ2H-Werte von Blattwachs n-Alkanen bisher wenig verstanden. Vor allen Dingen ist unklar, warum die n-Alkane in Gräsern, verglichen zu dikotylen Pflanzen, grundsätzlich in 2H abgereichert sind, und warum C3 Gräser stärker abgereichert sind als C4 Gräser. Auch ist der Zeitpunkt der n-Alkan Synthese und der de novo Synthese von n-Alkanen in voll entwickelten Blättern immer noch ein Diskussionsgegenstand. Bisher veröffentlichte Kalibrationsstudien, welche die Ursachen der Heterogenität von n-Alkan δ2H-Werten untersuchten, fokussierten dabei ausschliesslich auf die n-Alkane von Blättern. Für eine genauere Erforschung dieses Themas, wurden in dieser Doktorarbeit drei Studien durchgeführt, welche die Effekte von physiologischen und biochemischen Prozessen auf die δ2H-Werte von Blattwachs n-Alkanen in Gräsern untersuchen. In der ersten Studie wurde der Effekt der evaporativen Blattwasser-2H-Anreicherung (LW Δ2H) auf die δ2H-Werte der n-Alkane für eine Auswahl von C3 und C4 Gräsern bestimmt. Dafür wurden die Gräser unter kontrollierten Umweltbedingungen in Klimakammern kultiviert. Die Studie ergab, dass die LW Δ2H-Anreicherung nur teilweise in den δ2H-Werten der n-Alkane der C3 und C4 Gräser sichtbar ist. Darüber hinaus wurde auch gezeigt, dass sich die biosynthetische Fraktionierung der Wasserstoffisotopen (εbio) in C3 und C4 Pflanzen unterscheidet. Aus 6 Zusammenfassung diesem Grund wird angenommen, dass die δ2H-Werte der Blattwachs n-Alkane von C3 und C4 Gräsern nicht durch die LW Δ2H-Anreicherung bestimmt werden, sondern vielmehr durch systematische Unterschiede dieser beiden Pflanzengruppen in ihrer εbio. In der zweiten Studie lag der Fokus auf der Bestimmung des Zeitpunktes der Blattwachs n-Alkan Synthese in Gräsern. Dafür wurde die sekundäre Blattwachs n-Alkan Synthese in voll entwickelten Blattspreiten von C3 Gräsern untersucht. Dabei konnten die Experimente zeigen, dass Wasserstoff aus stark 2H-angereichertem Bewässerungswasser in Wachs n-Alkane von voll entwickelten Blättern eingebunden wurde. Dabei fand die Wachs n- Alkan Synthese in voll entwickelten Blättern auch nach dem Blattaustrieb weiterhin statt. Die Rate der sekundären n-Alkan Synthese war jedoch relativ tief und variierte für verschiedene Arten zwischen 0.09 und 1.09% pro Tag. Aufgrund dieser tiefen Rate werden die δ2H-Werte der Blattwachs n-Alkane vor allem durch die Umweltbedingungen in der Austriebsphase der Blätter festgelegt. In der dritten Studie wurden die n-Alkan Konzentrationen und die δ2H-Werte von verschiedenen Grasorganen untersucht. Dafür wurden Blätter, Blattscheiden, Stängel, Blütenstände und Wurzeln von 15
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