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The Fate of Recently Assimilated Carbon in Mature Deciduous Forest Trees

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The Fate of Recently Assimilated Carbon in Mature Deciduous Forest Trees The fate of recently assimilated carbon in mature deciduous forest trees Inauguraldissertation zur Erlangung der Würde eines Doktors der Philosophie vorgelegt der Philosophisch-Naturwissenschaftlichen Fakultät der Universität Basel von SONJA GISELA YIN KEEL aus Basel Basel, 2006 Genehmigt von der Philosophisch-Naturwissenschaftlichen Fakultät auf Antrag von Prof. Dr. Christian Körner Prof. Dr. Nina Buchmann Basel, den 19. September 2006 Prof. Dr. Hans-Jakob Wirz Dekan Acknowledgements First of all I would like to thank Rolf Siegwolf who offered me this PhD position. He introduced me to the fascinating world of stable isotopes and I learned a lot from him which went far beyond the topic of my thesis. We had a lot of fruitful and lively discussions which inspired my work and I felt extremely comfortable in his group. Many thanks to Christian Körner for giving me the opportunity to work in the forest canopy. He is an excellent teacher and I learned a lot about how to communicate science. I strongly appreciated his critical and very honest comments, his support and encouragement. I thank Nina Buchmann for her spontaneous and immediate positive response to co-examine my thesis. Thanks to Urs Baltensperger for investing a lot in a good team spirit. Although I, as a biologist, was an outsider in the Laboratory of Atmospheric Chemistry I felt as part of the group and even developed some interest for things that happen above the canopy. Many thanks to… Erwin Amstutz who manoeuvred me safely through the crowns, was extremely helpful and entertaining. Olivier Bignucolo who looked after the ‘isometers’ and collected leaf and litter samples. Georges Grun and Alain Studer for taking care of the web-FACE. Maya Jäggi for her great support in the lab. I very much enjoyed working with her. Steeve Pepin for his support and the good collaboration. He wrote me the most encouraging emails.Matthias Saurer for his assistance in the lab and for solving problems with ‘Analyza’ & Co. Katharina Steinmann for providing me data and answering a lot of questions. Markus Wilhelm, Thomas Brodtbeck, Kurt Minder and all other members of the ‘Pilzverein’ Basel who helped me with the taxonomic classification of the fungi. Erika Hiltbrunner who could answer nearly all my questions and gave me a hand in urgent situations. Günter Hoch for the good cooperation. Writing a paper with him was a real pleasure. Sebastian Leuzinger for statistical advice and for taking care of the weather data. Christina Reynolds, Christina Schädel & Cornelia Garbe for the extremely nice ambience in both my offices. Roman Asshoff for making me laugh. I miss his humour… Susi Peláez-Riedl, the ‘master of graphs and slides’ for helpful suggestions. Rossella Guerrieri supported the sample preparation and was a most pleasant guest. René Richter for help in case of technical emergency. Pascal Niklaus for introducing me to ‘R’ and for statistical advice. Franziska Grob for her great efficiency and friendliness. Doris Hirsch-Hoffmann for being so well organized. Kristin Strobel who collected part of the aphids. I am thankful to Hafdis-Hanna Ægisdóttir, Martin Bader, Cornelia Garbe, Tanya Handa, Sebastian Leuzinger, José Monteiro, Katrin Rudmann, Christina Schädel, Eva Spehn, Stefan van Ekeren for their special support during the last few weeks of my PhD. Also Isabelle Frede took great care of me. I am grateful for her long lasting friendship. I very much appreciated the familiar and friendly atmosphere at the Institute of Botany at the University of Basel as well as in the Laboratory of Atmospheric Chemistry at the Paul Scherrer Institute and would like to thank all other members, too. I would like to thank my parents Ivy and Peter Keel, my sister Jenni and Elias Wieland for their love and support through all the years. Unfortunately my grandparents Agnes and Albert Keel could only witness the beginning of my PhD. They always showed great interest in my work. I thank my grandmother Louise Cheng who came all the way from Canada for my defence. Last but not least I would like to thank Stephan Schaffner for his great love. During the last weeks of my PhD, he gave me the necessary strength and could cheer me up when I was desperate. He spent hours quietly studying Swedish next to me, while I was working and I very much enjoyed his company. Table of contents Chapter 1 General introduction 7 Chapter 2 Canopy CO2 enrichment permits tracing the fate of recently 11 assimilated carbon in a mature deciduous forest Chapter 3 Strict separation of carbon transfer to mycorrhizal and 23 saprophytic fungi by tall temperate forest trees Co-authorship in Siegwolf et al., in revision Chapter 4 Carbon flux and growth in mature deciduous forest 33 trees exposed to elevated CO2 Co-authorship in Körner et al. (2005) Chapter 5 Tracing arthropod movement in a deciduous forest 37 canopy using stable isotopes Co-authorship in Asshoff et al. (2006) Chapter 6 Rapid mixing between old and new carbon pools in the 47 canopy of mature forest trees Chapter 7 13C-labelling reveals different contributions of photo- 57 assimilates from infructescences for fruiting in two temperate forest tree species Co-authorship in Hoch & Keel (2006) Chapter 8 Stomatal conductance of mature forest trees exposed to 67 elevated CO2 Chapter 9 General summary 77 Curriculum vitae 81 5 Chapter 1 General introduction This thesis made use of a specially designed free-air CO2 enrichment (FACE) system which was set up in a mature Forests store half of the terrestrial organic carbon (C) of deciduous forest near Basel in the year 2000. The so which two-thirds are contained in soils (Dixon et al., called web-FACE (Pepin & Körner, 2002) complies with 1994). Nearly all biomass C is assimilated by leaves from the heterogeneous structure of a deciduous forests' where C-compounds are transferred to carbon sinks such canopy and its height of 30-36 m. Due to its fossil fuel as growth and reproduction, maintenance, storage or origin the stable isotope ratio of the supplemental CO2 export (Körner, 2003). How rapidly recently assimilated differs from atmospheric CO2, leaving a distinct signal in C is transferred to these various C pools and fluxes is so all compartments where C enters. Potted C4 grasses far largely unknown, in particular it is not well grown in the tree crowns served as a reference for the understood at which speed C is allocated to long-lived isotope signals (referred to as ‘isometers’). Because the pools. Depending on where C is allocated to, it stays in grasses consist exclusively of C that originates from the CO2 they assimilate, with no influence from old C the system for minutes (e.g. leaf respiration) to hundreds 13 or thousands of years (e.g. recalcitrant soil C). reserves, the δ C difference of grasses exposed to Within trees assimilated C is not evenly distributed labelled air compared with grasses grown in ambient air due to different sink strengths. Carbon is directed to areas reflects the actual isotopic signal the canopy is exposed of most active growth which strongly vary through the to. Using a rule of proportion where the isometer signal season. For instance the often observed bimodal growth of 5.9‰ refers to 100% new C, the fraction of labelled C of fine roots, with a first peak in early spring and a which had reached every specific forest compartment second in autumn (Shiroya et al., 1966; Smith & Paul, could be calculated. 1988), is the result of a fine tuned seasonal change in C Compared to the conventional FACE systems the allocation within the tree. web-FACE enriches the tree crowns only. Since the Indirect measurements based on correlations of the canopy is high enough, no labelled CO2 reaches the forest floor by direct diffusion or downward draughts as was isotopic composition of soil respired CO2 with climatic conditions of previous days indicate that CO is respired confirmed by stable isotope analysis of understory herbs 2 13 within only 4-10 days after assimilation (Ekblad & showing no C signals (Steinmann et al., 2004). This Högberg, 2001; Bowling et al., 2002; Steinmann et al., offers to trace the pathway of recently assimilated C 2004; Knohl et al., 2005). Roughly half of the C through stems into roots, soil and soil air without assimilated by trees, is rapidly returned to the atmosphere confounding CO2 uptake via understory vegetation or (Högberg et al., 2002) by above ground respiration direct diffusion. (Horwath et al., 1994; Nogués et al., 2006) as well as Although this large scale C allocation study was below ground respiration (Andrews et al., 1999; Högberg carried out within a CO2 enrichment experiment, the et al., 2001; Andersen et al., 2005). To assess residence main focus here is not on CO2 effects as such. This thesis times of pools that are too young for conventional rather capitalizes on the unavoidable C-labelling 14 associated with CO2 enrichment. We assume that radiocarbon dating the so called ‘bomb- CO2’ peak, which is the result of nuclear weapons tests during the elevated CO2 did not have any major consequences for early 1960ies can be used as a reference. Using this the speed and direction of C flows or C pools. Under this method fine roots were found to be 3-18 years old assumption the data provide an unique opportunity for (Gaudinski et al., 2001) and more than ten year old C assessing the fate of carbon in situ in a tall forest. could be detected in saprophytic fungi (Hobbie et al., In the first year of the canopy labelling a striking 2002). Longest lived pools are found in soil organic species-specificity in the amount of new C which had matter reaching ages up to several thousand years been incorporated into leaves was found (Steinmann et (Trumbore, 2000).
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