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The Chemistry of Specificity in the Rinorea- Cymothoe Host-Herbivore

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The Chemistry of Specificity in the Rinorea- Cymothoe Host-Herbivore THE CHEMISTRY OF SPECIFICITY IN THE RINOREA - CYMOTHOE HOST -HERBIVORE ASSOCIATION AN EXPERIMENTAL INVESTIGATING INTO THE CHEMICAL BASIS FOR RECOGNITION BY THE AFROTROPICAL SPECIALIST HERBIVORE CYMOTHOE EGESTA (N YMPHALIDAE ), OF ITS HOST PLANT RINOREA ILICIFOLIA (V IOLACEAE ) AUTHOR : MS C THESIS GERCO S. NIEZING AUGUST 2011 REG . NO. 860414 604 010 BIS-80436 SUPERVISION : FREEK T. BAKKER -BIOSYSTEMATICS GROUP ROBIN VAN VELZEN -BIOSYSTEMATICS GROUP MAARTEN A. JONGSMA -PRI BIOSCIENCE THE CHEMISTRY OF SPECIFICITY IN THE RINOREA -CYMOTHOE HOST -HERBIVORE ASSOCIATION NIEZING , AUGUST 2011 2 THE CHEMISTRY OF SPECIFICITY IN THE RINOREA -CYMOTHOE HOST -HERBIVORE ASSOCIATION NIEZING , AUGUST 2011 Title Page: 1: View of Kakum National Park from the Canopy Walkway 1 2 2: Cymothoe egesta eggs on a Rinorea ilicifolia leaf 3: Cage experiment with a female Cymothoe egesta feeding on coconut 3 4 4: Cymothoe egesta caterpillar on a young Rinorea ilicifolia leaf 5: LC-MS chromatogram, i.e. “the chemistry underlying host specificity” 5 1 THE CHEMISTRY OF SPECIFICITY IN THE RINOREA - CYMOTHOE HOST -HERBIVORE ASSOCIATION AN EXPERIMENTAL INVESTIGATING INTO THE CHEMICAL BASIS FOR RECOGNITION BY THE AFROTROPICAL SPECIALIST HERBIVORE CYMOTHOE EGESTA (N YMPHALIDAE ), OF ITS HOST PLANT RINOREA ILICIFOLIA (V IOLACEAE ) AUGUST 2011 MS C THESIS BIS-80436 Author: Gerco S. Niezing, BSc Reg. No. 860414 604 010 [email protected] Supervisors: Dr. Freek T. Bakker Biosystematics Group [email protected] Ir. Robin van Velzen Biosystematics Group [email protected] Dr. Ir. Maarten A. Jongsma PRI Bioscience [email protected] 3 THE CHEMISTRY OF SPECIFICITY IN THE RINOREA -CYMOTHOE HOST -HERBIVORE ASSOCIATION NIEZING , AUGUST 2011 PREFACE It was my ambition to have this thesis do two things for me personally: allow me to go somewhere far, warm and adventurous, as well as let me have a taste of scientific research. The first goal was met ten times over. I lived with a local family in a Ghanaian village where I connected intimately with the local culture and – every biologist’s dream – I could run around in a tropical rainforest. With regard to the second goal, I have explored some very diverse scientific aspects: field work, behavioural experimentation as well as lab work with cutting edge chemical analyses. At the top-level, this thesis deals with macroevolution, the science of speciation and the basis of biodiversity; this fascinates me more than anything. The methods I have used have never been performed in this manner before to my knowledge and that in my opinion already warrants this study. I did however often wonder whether I had not bitten off more than I could chew in constructing this project. Whilst actually being an animal biologist, I have herewith performed an investigation in the fields of ecology and phytochemistry using experimental methods that have never been done in this way before, at a chair group that normally deals with phylogenetic studies. Still, I am very well aware of the fact that the MSc thesis investigation is a training period. I have learned a great deal about science, evolution, the world and myself in the past year. It is my only hope that the esteemed reader will also learn something about the interesting world of insect-plant biology and the role of chemistry in its evolution. Gerco Niezing Wageningen, August 2011 4 THE CHEMISTRY OF SPECIFICITY IN THE RINOREA -CYMOTHOE HOST -HERBIVORE ASSOCIATION NIEZING , AUGUST 2011 TABLE OF CONTENTS Preface ....................................................................................................... 4 Table of Contents ........................................................................................ 5 Summary ................................................................................................... 6 Chapter 1 Introduction ............................................................................... 7 1.1 General introduction ......................................................................... 7 1.2 The case of Rinorea and Cymothoe ....................................................10 1.3 Oviposition preference as a proximate cause of host specificity .............12 1.4 Identifying chemicals involved in host recognition by Cymothoe ............13 1.5 Identifying chemicals involved in host recognition by Cymothoe ............14 1.6 Model association Rinorea ilicifolia-Cymothoe egesta ...........................16 1.7 Untargeted metabolomics approach ...................................................18 1.8 Testing of CPG as oviposition stimulant ..............................................23 1.9 Aims and hypotheses .......................................................................24 Chapter 2 Field work: set-up and observations .............................................25 2.1 Set-up and location .........................................................................25 2.2 Observations of Cymothoe egesta (oviposition) behavior ......................27 2.3 Miscellaneous observations ...............................................................32 Chapter 3 Untargeted metabolomics ...........................................................34 3.1 Method and materials ......................................................................34 Procedure in field work and sampling protocol ...........................................34 Metabolite extraction ..............................................................................35 LC PDA QTOF MS analysis .......................................................................36 Data processing .....................................................................................37 Metabolite identification ..........................................................................39 3.2 Results ..........................................................................................40 Sampling of Rinorea ilicifolia plant material ...............................................40 LC PDA TOF MS analysis and data processing ............................................41 Data analysis and peak selection .............................................................42 Metabolite identification ..........................................................................52 Discussion ............................................................................................54 Chapter 4 Testing of CPG as oviposition stimulant ........................................55 4.1 Obtaining 2-(3’-cyclopentenyl)glycine ................................................55 4.2 Testing for the presence of CPG using LC-MS ......................................57 Method .................................................................................................57 Results .................................................................................................58 Discussion ............................................................................................59 4.3 Behavioral experiment .....................................................................61 Methods and materials ...........................................................................61 Results and Discussion ...........................................................................65 Chapter 5 Conclusions ...............................................................................68 Chapter 6 Recommendations .....................................................................69 Acknowledgements .....................................................................................70 References ................................................................................................71 Appendix Preliminary testing of R. yaundensis .............................................76 5 THE CHEMISTRY OF SPECIFICITY IN THE RINOREA -CYMOTHOE HOST -HERBIVORE ASSOCIATION NIEZING , AUGUST 2011 SUMMARY Plants and phytophagous insects together show an enormous diversity. Studying the driving factors behind the diversification of these groups is a major topic in ecology and evolutionary biology. One of the most striking observations regarding this topic is the high level of host specialization by herbivorous insects. There may be a process of reciprocal co-evolution at the basis of the observed patterns. Most evidence for this however, is gathered with clear cases of plant defence and insect adaptation to these defences. Whether plants radiate because of selective pressure from insects, and what causes insects to be host specific even on apparently undefended plants is still not quite understood. This MSc thesis deals with these issues using the model host-herbivore association Rinorea-Cymothoe . This is an “extreme case” of host specificity involving a large number of congeneric species, and without clear co-adaptation regarding (anti-)herbivory. Within the WU biosystematics group, this association is being study by PhD student Robin van Velzen, who is performing phylogenetic studies of both genera to gain insight into the evolutionary patterns. In order to make this a more integrative study on different levels, I have attempted to uncover the chemical basis for the proximate cause of host specificity: female oviposition preference. To this end, I have conducted field work in Kakum National Park, Ghana, during which I observed and experimented on the model association Rinorea ilicifolia-Cymothoe egesta . I had two main approaches: 1) untargeted metabolomics to select hypothetical oviposition stimulants to Cymothoe egesta , and 2)
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