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02 Whole.Pdf (3.548Mb) Copyright is owned by the Author of the thesis. Permission is given for a copy to be downloaded by an individual for the purpose of research and private study only. The thesis may not be reproduced elsewhere without the permission of the Author. The Physiology and Control of Re-greening in Spathes of Zantedeschia A thesis presented in partial fulfilment of the requirements for the degree of Doctor of Philosophy in Plant Physiology at Massey University, Palmerston North, New Zealand Jianyu Chen 2011 Abstract Abstract Spathe re-greening is a primary determinant limiting postharvest quality of Zantedeschia Spreng. as a cut flower, pot- or landscape-plant. A treatment that can be utilised by growers to delay re-greening offers potential to improve the postharvest quality and provide a marketing advantage. To achieve this, and develop an understanding of the physiological mechanism of re-greening, this project investigated the changes in colour, levels and types of pigment, and differentiation of plastids in spathe tissue durng re-greening; and how this process was controlled by various factors including fructification, light and various plant hormones (e.g. cytokinin and gibberellin). In the hybrid ‘Best Gold’, spathe re-greening was initiated within three days after horticultural harvest-maturity and, within two weeks the whole abaxial surface of the spathe had re-greened. During this period, the adaxial surface did not re-green and remained yellow in colour. The change in colour of the abaxial surface primarily resulted from the accumulation of chlorophyll within the subepidermal layers, as reflected by a strong correlation between the colour coordinate hue angle (Hº) and total chlorophyll content in that surface (r = 0.98). Monitoring Hº can therefore, be used to evaluate the degree of re-greening for ‘Best Gold’ without chlorophyll analysis. The content of carotenoid (in particular lutein which was predominant) was comparatively steady during re-greening. From an ultrastructural perspective, spathe re-greening was characterized by redifferentiation of chloroplasts from chromoplasts, as compared with de novo synthesis of chloroplasts from proplastids. The redifferentiation of chloroplasts involved thylakoid reformation through multiple i Abstract mechanisms. In addition to de novo synthesis of thylakoid by invaginations of the inner-envelope membrane, it is likely that the thylakoids were either derived from primary thylakoids or plastoglobuli present in mature chromoplasts. The occurrence of re-greening in the spathe of both Zantedeschia aethiopica and ‘Best Gold’ following the removal of the spadix prior to pollination, contradicted the hypothesis that re-greening was induced by fructification (Pais and Neves, 1982- 1983). Further to this, the occurrence of re-greening in the spathe of ‘Best Gold’ with a spadix naturally devoid of female flowers, and the re-greening of pigmented leaves devoid of any true flower parts, also contradicted this hypothesis. The current findings therefore, indicate that fructification is not necessarily a prerequisite for induction of re-greening. In absence of light, no chlorophyll accumulated in spathe tissue of ‘Best Gold’, but the initial redifferentiation of chloroplasts from chromoplasts, as characterized by the formation of double-membrane lamella, was noted. Without light however, redifferentiation of chloroplasts was not completed. These suggest spathe re-greening requires light for the process to complete, but the onset of re-greening can be induced in darkness. The application of 6-benzylaminopurine (BAP) stimulated re-greening in spathe tissue by enhancing accumulation of carotenoid and chlorophyll, and also stacking of grana. But the response to BAP was dependent on the presence/absence of light, the stage of re-greening, and which surface, i.e. abaxial or adaxial. In contrast, the application of gibberellin (GA3) retarded formation of double-membrane lamella, and thus delayed the onset of re-greening. Hence, a synergistic effect of BAP and GA3 in delaying the onset of re-greening was likely to be a result of co-regulation between ii Abstract BAP-stimulated accumulation of carotenoid and GA3-stimulated retardation of chloroplast redifferentiation. By integrating both light and hormonal factors, several methods were tested on the actual horticultural commodity, i.e. flowers (peduncle, spathe and spadix) of ‘Best Gold’, so as to evaluate their efficacy in delaying re-greening. Pulsing flowers in darkness at 5 ºC for 24-h in a solution containing both GA3 and BAP, was the most effective treatment in postponing re-greening, resulting in a seven-day delay in re- greening of the spathe. iii Acknowledgements Acknowledgements I offer my deepest gratitude to my chief supervisor Dr. Keith Funnell. Without your excellent guidance, encouragement, patience, and support throughout this study, completion of this thesis would have not been possible. Your passion for science has made this journey through my PhD very enjoyable. Your dedication and time in reading and critically evaluating my thesis from pieces at the very beginning to final drafts is also very much appreciated. My special gratitude also goes to my co-supervisor Dr. David Lewis. Thanks for your invaluable advice and help whenever I needed it for my thesis. Your feedback on my thesis was very important to complete this project. Your challenges to my interpretation of the results have inspired me to consider many new ways of thinking. Your classic words of encouragement e.g., “hang in there” have kept me going during the hard times of writing. I would like to thank Dr. Jocelyn Eason for co-supervising this thesis. You have always believed in my abilities and provided guidance and encouragement whenever I needed it. Your efficiency at providing feedback on chapters was legendary given your demanding workload. I would like to express my gratitude to Dr. David Woolley for co-supervising this thesis. Your expertise and enthusiasm in the areas of plant hormones and chromatography have in part enabled the success of this project. Your feedback on the thesis is valuable and very much appreciated. iv Acknowledgements I am indebted to many staff members within the Institute of Natural Resources, Massey University for their help throughout this project. In particular, to Mr James Slater, thank you for your friendship and excellent assistance in many ways including growing plants, collecting samples and pigment analysis, even after working many hours. To the staff at Plant & Food Research Ltd., particularly Steve Arathoon, thank you for your technical assistance in pigment extraction. To my colleagues at the Manawatu Microscopy & Imaging Centre: Mr. Doug Hopcroft, thank you for your instruction and assistance in both light and electron microscopy, for your support, laughter, and jokes, and for teaching me those ‘essential’ life skills (e.g. speaking in Kiwi slang); To Dr. Dmitry Sokolov, thanks for your technical assistance in Confocal Microscopy; To Dr. Al Rowland, thank you for reading my thesis and providing me with valuable suggestions; To Professor Barry Scott, thank you for your support and allowing me to take time off from work to write the thesis; To Ms Pat Munro, thank you for your care, kindness and big smiles, which always brighten my day. I am also grateful to Mr. Jim Wilson and Golden State Bulb Growers for providing plant material; The Sir Victor Davies Foundation for partial funding of this research; Mr Jim Wilson for partial financial assistance for me personally during the latter parts of this research; Education NZ for providing an International Doctoral Research Scholarship for my study. My special appreciation is also extended to my friends in NZ: to Maggie Zou for your awesome friendship, encouragement and yummy food; To the Phoa family: v Acknowledgements Liyan, Patrick, Auntie Kong, Uncle Li and little Emmanuel, thanks for all the joy, blessings and making me feel at home. Finally, I owe so much to my parents and brother for their love, patience, and support while I was far away from them. Ba and Ma, Gege, I love you all very much. vi Table of Contents Table of Contents ABSTRACT ................................................................................................................................ I ACKNOWLEDGEMENTS ..................................................................................................... IV TABLE OF CONTENTS ...................................................................................................... VII LIST OF FIGURES .............................................................................................................. XIII LIST OF TABLES .................................................................................................................. XX LIST OF ABBREVIATIONS .............................................................................................. XXI CHAPTER 1 GENERAL INTRODUCTION .................................................................... 1 1.1 Zantedeschia .............................................................................................................. 1 1.2 Spathe re‐greening of Zantedeschia ............................................................................ 5 1.3 Re‐greening in organs of other plant species ............................................................... 8 1.4 Greening, de‐greening, re‐greening and senescence.................................................. 11 1.4.1 Plastids ..................................................................................................................
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