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Fern Histology: a Focus on Cell Walls and Aspleniaceae

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Fern Histology: a Focus on Cell Walls and Aspleniaceae . I could exceedingly plainly perceive it to be all perforated and porous, much like a Honey- comb, but that the pores of it were not regular . these pores, or cells, . were indeed the first microscopical pores I ever saw, and perhaps, that were ever seen, for I had not met with any Writer or Person, that had made any mention of them before this . Drawing of dried cork tissue 1 Robert Hooke coined the term ‘cell’ for describing units in plant tissues. 1 Robert Hooke (1665) Micrographia: or, some physiological descriptions of minute bodies made by magnifying glasses. J. Martyn and J. Allestry, London. Supervisor/Promotor: Prof. Dr. Ronnie Viane (University of Ghent) Members of the jury/Leden van de jury: Prof. Dr. Luc Lens (University of Ghent, chairman) Prof. Dr. Ronnie Viane (University of Ghent) Prof. Dr. Paul Knox (University of Leeds) Prof. Dr. Tom Beeckman (University of Ghent/Flanders Institute for biotechnology) Prof. Dr. Olivier De Clerck (University of Ghent) Prof. Dr. Annemieke Verbeken (University of Ghent) Date of Public defence : 20 th May 2011 This PhD research was performed in the Research Group Pteridology, Department of Biology, Faculty of Sciences, Ghent University, K.L. Ledeganckstraat, B-9000 Gent. Dit doctoraatsonderzoek werd uitgevoerd in de Onderzoeksgroep Pteridologie, Vakgroep Biologie, Faculteit Wetenschappen, Universiteit Gent, K.L. Ledeganckstraat, B- 9000 Gent. Correct citation: Leroux O. 2011. Fern histology: a focus on cell walls and Aspleniaceae. PhD-dissertation. Ghent University, Belgium. FERN HISTOLOGY: A FOCUS ON CELL WALLS AND ASPLENIACEAE VAREN HISTOLOGIE: EEN FOCUS OP CELWANDEN EN ASPLENIACEAE Olivier LEROUX Thesis submitted in fulfillment of the requirements for the degree of Doctor (PhD) in Sciences (Biology) Proefschrift ingediend tot het behalen van de graad van Doctor in de Wetenschappen (Biologie) Academic year 2010-2011 Academiejaar 2010-2011 Acknowledgements From experience I can tell you that these few pages are among the most widely read pages of most dissertations. It is here where you think that you will find out whether or not you have meant something in the life of a PhD student. While this may be true to some extent, to me, it seems an impossible task to mention all the people that have crossed my professional path during the last six years. I am going to try anyway and if your name is not listed, rest assured that my gratitude is not less than for those listed below. First and foremost, this thesis would not have been possible without the help, support and patience of my supervisor, Ronnie Viane. Thanks for encouraging me to develop my own ideas and, especially, for teaching me the skills of microscopic observation and microphotography. Most of the work presented here would have been impossible without the generous support, advice and guidance of Paul Knox. Thank you for giving me the opportunity to carry out some of the practical work in your lab and also for introducing me in the fascinating world of cell wall research. It is also a pleasure to pay tribute to the many collaborators and co-authors. In particular, I would like to thank Agnieszka Bagniewska-Zadworna (for helping me with scanning electron microscopy, chapter 1, 3, 5, 6, 9, and 10), Siti Khadijah Rambe and Dirk Stubbe (for the molecular analysis, chapter 4), Antonio Pizzi, Brigitte Chabbert and Anouk Habrant (for the lignin analyses, chapter 4 and 5), and Sue Marcus (for providing some immunolabelling micrographs, chapter 8 and 9). I am most grateful to Guy Van Der Kinderen, for sharing a lab/office, and, unintentionally, my (occasional) messiness. Your technical experience and your help with realizing some of the ideas in my quest for cheaper alternatives in science are most appreciated. I also wish to thank William Willats and Iben Sørensen (for the CoMPP-analysis, chapter 7), and Zoë Popper (for the interesting discussions on different aspects of this thesis). I was fortunate in having had the advice and help from two experienced electron microscopists. Riet and Myriam, thank you for answering my many questions. I would also like to thank my twin brother Frederic. Next to sharing nearly identical DNA, we also share the passion for microscopy and this mutual interest resulted in some nice TEM-images in chapter 4 and 6. I am thankful to Tom Beeckman, Annemieke Verbeken, and Olivier De Clerck. In the midst of all their activity they accepted to be members of the examination committee. I acknowledge their valuable comments and suggestions on the draft version of this thesis. I also would like to express my gratitude to Dominique Vanderstraeten, Paul Goetghebeur, and Marc Espeel. I was fortunate to have had the opportunity to collaborate with the Centre for X-ray Tomography (UGCT). My sincere thanks are due to Bert, Denis, and Manuel, for scanning many plant samples. I also wish to thank all my colleagues of the Ledeganck building (Botany, Physiology, and Zoology research groups) for enduring my presence: Adelin, Barbara, Bram, Carolina, Dieter, Dirk R, Dirk S, Elien, Elke, Eske, Filip, Francine, Isabel, Jan, Janina, Jasper, Joke, Jorinde, Kobeke, Kristof, Leander, Liesbeth, Marc, Marie-Stéphanie, Marjolein, Marleen, Peter, Pieter, Rosette, Wim, Yannick, ... As I do not wish to embarrass anybody with some quite interesting anecdotes, I decided not to include any here. I’m grateful to the staff of the botanical garden for cultivating many of the plants I studied and often killed (for the sake of science): Anneke, Chantal, Herbert, Jean-Pierre, Marc D, Marc L, Olivier, Ritchy, Stephan, Stijn and Thierry. I was fortunate to have spent many weeks at the Knox Lab and I would like to thank everybody in Leeds for making every stay so enjoyable: Alenka, Carem, Cécile, Hannah, José, Kieran, Li, Sue, Thomas, Tom, Tony and Yves. I would also like to acknowledge the (so much needed) financial support from the Fund for Scientific Research —Flanders (F.W.O. — Vlaanderen) and the Systematics Research Fund (The Linnean Society of London). I’m also indebted to my parents for their unparalleled interest and encouragement, and to my friends (I think I managed not to talk about work that often). Last but not least, I would like to thank Mother Nature for making plants (and cell walls in particular) so excitingly and wonderfully complex. (Perhaps, I forgot someone … so, just in case: thank you to whom it concerns!) Table of contents PART 1: INTRODUCTION & TECHNICAL PAPERS CHAPTER 1: GENERAL INTRODUCTION ....................................................................... 1 1.1 HISTORY OF (FERN) ANATOMY: A BRIEF SUMMARY ........................................ 1 1.2 FERNS AND LYCOPHYTES ......................................................................................... 2 1.3 ASPLENIACEAE ............................................................................................................. 4 1.4 PLANT CELL WALLS .................................................................................................... 7 1.5 CELL WALLS OF FERNS AND LYCOPHYTES ....................................................... 10 1.6 GENERAL AIMS AND OUTLINE .............................................................................. 12 1.7 REFERENCES ................................................................................................................ 15 CHAPTER 2: A SANDWICH-EMBEDDING METHOD FOR ORIENTED SECTIONING ......................................................................................................................... 25 2.1 INTRODUCTION .......................................................................................................... 26 2.2 MATERIALS AND METHODS ................................................................................... 27 2.3 RESULTS AND DISCUSSION ..................................................................................... 30 2.4 CONCLUSIONS ............................................................................................................ 32 2.5 REFERENCES ................................................................................................................ 32 CHAPTER 3: A NEW PREPARATION METHOD FOR STUDYING FRESH PLANT STRUCTURES USING X-RAY COMPUTED TOMOGRAPHY .................................... 35 3.1 INTRODUCTION .......................................................................................................... 36 3.2 MATERIAL AND METHODS ..................................................................................... 37 3.3 RESULTS AND DISCUSSION ..................................................................................... 39 3.4 CONCLUSIONS ............................................................................................................ 43 3.5 REFERENCES ................................................................................................................ 43 PART 2: ASPLENIACEAE: ROOT ANATOMICAL FEATURES CHAPTER 4: HISTOGENY, STRUCTURE, COMPOSITION AND TAXONOMICAL SIGNIFICANCE OF THE SCLERENCHYMA SHEATH IN ASPLENIACEAE ROOTS (POLYPODIALES)................................................................................................... 51 4.1 INTRODUCTION .......................................................................................................... 52 4.2 MATERIALS AND METHODS ................................................................................... 54 4.3 RESULTS ........................................................................................................................
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