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Floral Scent in Salix L. and the Role of Olfactory and Visual Cues for Pollinator Attraction of Salix Caprea L

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Floral Scent in Salix L. and the Role of Olfactory and Visual Cues for Pollinator Attraction of Salix Caprea L Floral Scent in Salix L. and the Role of Olfactory and Visual Cues for Pollinator Attraction of Salix caprea L. Dissertation zur Erlangung des Doktorgrades Vorgelegt der Fakultät für Biologie, Chemie und Geowissenschaften der Universität Bayreuth von Ulrike Füssel Bayreuth, im Oktober 2007 II Die Arbeit wurde von August 2004 bis Oktober 2007 am Ökologisch-Botanischen Garten der Universität Bayreuth in der Arbeitsgruppe von Herrn PD Dr. Gregor Aas angefertigt. Gefördert wurde die vorliegende Arbeit durch ein Stipendium der Deutschen Forschungsgemeinschaft (Graduiertenkolleg 678 – Ökologische Bedeutung von Wirk- und Signalstoffen bei Insekten – von der Struktur zur Funktion). Vollständiger Abdruck der von der Fakultät für Biologie, Chemie und Geowissenschaften der Universität genehmigten Disseration zur Erlangung des Grades eines Doktors der Naturwissenschaften (Dr. rer. nat.). Tag der Einreichung: 24. Oktober 2007 Tag des Kolloquiums: 09. Januar 2008 Prüfungsausschuss PD Dr. G. Aas (Erstgutachter) Prof. Dr. K. H. Hoffmann (Zweitgutachter) Prof. Dr. K. Dettner (Vorsitzender) Prof. Dr. S. Liede-Schumann Prof. Dr. R. Schobert III This dissertation is submitted as a “Cumulative Thesis“ that includes four (4) publications: two (2) published articles, one (1) submitted article, and one (1) article in preparation for submission. The publications are listed in detail below. Published: • Dötterl S., Füssel U., Jürgens A., and Aas G. (2005): 1,4-Dimethoxybenzene, a floral scent compound in willows that attracts an oligolectic bee. Journal of Chemical Ecology 31:2993-2998 (Part B, Chapter 3). • Füssel U., Dötterl S., Jürgens A., and Aas G. (2007): Inter- and intraspecific variation in floral scent in the genus Salix and its implication for pollination. Journal of Chemical Ecology 33:749-765 (Part B, Chapter 1). Submitted: • Füssel U., Dötterl S., Jürgens A., Woodring J., and Aas G. (2008): Floral reward and advertisement in dioecious Salix caprea. Submitted to Plant Biology (Part B, Chapter 4). Prepared for resubmission: • Füssel U., Dötterl S., Jürgens A., and Aas G. (2008): Salix caprea: An interaction generalist and multi-specialist with bimodal adaptations of floral scent to bees and moths. Intended for resubmission to New Phytologist (Part B, Chapter 2). IV Declaration of Self-Contribution of Research The thesis contains a detailed summary (Part A) and four (4) research articles (Part B), covering various research work on pollination biology and chemical ecology of willows and their pollinators. Most of the research work presented in this thesis was carried out by myself independently at the Ecological-Botanical Garden, University Bayreuth under supervision of PD Dr. Gregor Aas. Together with my supervisor and all co-authors (Dr. Stefan Dötterl, Dr. Andreas Jürgens, and Prof. Dr. Joseph Woodring) I developed the methods, discussed the results and prepared the manuscripts of all research articles. My practical field and laboratory work was supported by several students and employees of the Ecological-Botanical Garden and the University. 1st article Füssel U., Dötterl S., Jürgens A., and Aas G. (2007): Inter- and intraspecific variation in floral scent in the genus Salix and its implication for pollination. Journal of Chemical Ecology 33:749-765. (Part B, Chapter 1) My contribution to this chapter was about 85 %. The experimental design, the main part of the field work as well as the analysis, the presentation, and the interpretation of the results were performed by myself. 2nd article Füssel U., Dötterl S., Jürgens A., and Aas G. (submitted 2007): Salix caprea: An interaction generalist and multi-specialist with bimodal adaptations of floral scent to bees and moths. Intended for resubmission to New Phytologist. (Part B, Chapter 2) My contribution to this article was approximately 75 %. Dr. Stefan Dötterl conducted the GC-EAD study and analysed the GC-EAD data. The floral scent samples needed for the electrophysiological measurements were collected and prepared by myself. All the other data were also collected, analysed, presented, and interpreted by myself. Susanne Kern helped to collect and identify flower visitors. V 3rd article Dötterl S., Füssel U., Jürgens A., and Aas G. (2005): 1,4-Dimethoxybenzene, a floral scent compound in willows that attracts an oligolectic bee. Journal of Chemical Ecology 31:2993-2998. (Part B, Chapter 3) My contribution to this study was circa 60 %. The data concerning the behavioural experiment with 1,4-dimethoxybenzene were collected, analysed, presented, and interpreted completely by myself. Floral scent samples for the electrophysiological measurements were collected and prepared completely by myself. Data of the electrophysiological measurements were collected, presented, interpreted and discussed by Dr. Stefan Dötterl. He wrote also the first manuscript draft. 4th article Füssel U., Dötterl S., Jürgens A., Woodring J., and Aas G (prepared for submission): Floral reward and advertisement in dioecious Salix caprea. Submitted to Plant Biology. (Part B, Chapter 4) My contribution to this manuscript was about 80 %. The experimental design, the main part of the work in field and laboratory, as well as the analysis, presentation, interpretation and discussion of the results were performed by myself. Prof. Dr. Joseph Woodring introduced me in the HPLC method and I performed the nectar analyses myself. VI Acknowledgements This work would not have been possible without the help of many people and I would like to express my gratitude to all of them. First of all, I want to thank my supervisor, PD Dr. Gregor Aas for his kind support and the opportunity to work at the Ecological-Botanical Garden, Bayreuth. The many fruitful discussions with him helped to work out the importance of essential results. I am grateful to Prof. Dr. Sigrid Liede-Schumann for the possibility to perform the GC-MS analyses in the laboratories of the Department of Plant Systematics. Dr. Stefan Dötterl’s enthusiasm for chemical ecology, pollination and statistical analyses inspired me to work in this area of research. Sophie Cralischeck, Susanne Kern, and Nadja Nikol I want to say thank you for their cooperation. I want to thank all present and former members of the Ecological-Botanical Garden for creating a good working atmosphere and helping in many ways. Further I thank Dr. Andreas Jürgens and Dr. Taina Witt for their helpful comments and discussions on earlier drafts of the manuscripts. I am thankful to Prof. Dr. Joseph Woodring for correction of English language and style. I also want to thank Dr. Andreas Reuter for helpful discussions as well as for practical help. Particularly I want to thank all the students who helped and supported my work. With their assistance it was possible to perform all the time-consuming experiments during the short flowering time of the willows. Thanks to all members of the Graduate College 678 for their good cooperation. My special thanks go to my family, especially to my parents and my sister for their continuous love and help. I want to express many thanks to Thorsten for his love, help, understanding and patience. VII This project was financed by the Deutsche Forschungsgemeinschaft (Graduate College 678: Ecological Significance of Natural Compounds and other Signals in Insects – from Structure to Function). 1 Content Content ................................................................................................................... 1 Part A Detailed Summary ...................................................................................................... 3 1 General Introduction and Aims of the Research ................................................................ 4 1.1 Background ................................................................................................................ 5 1.1.1 The Genus Salix: Distribution and Taxonomy................................................... 5 1.1.2 Pollination System of Salix ................................................................................ 5 1.1.3 Floral Signals and Rewards of Salix .................................................................. 6 1.2 Aims of the Research ............................................................................................... 11 2 Material and Methods....................................................................................................... 12 2.1 Plant Material........................................................................................................... 12 2.2 Determination of Flower Visitors (Publication 2).................................................... 12 2.3 Floral Scent Collection and Analysis (Publications 1, 2, and 4).............................. 13 2.4 Gas Chromatography Coupled to Electroantennographic Detection (GC-EAD) (Publications 2 and 3)............................................................................................... 14 2.5 Behavioural Tests (Publications 2, 3, and 4)............................................................ 16 2.6 Sugar Composition and Concentration of Nectar in Flowers of Salix caprea (Publication 4).......................................................................................................... 21 2.7 Pollination Experiment (Publication 2).................................................................... 21 3 Results and Discussion..................................................................................................... 22 3.1 What Is the Chemical Composition of Salix
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