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Evidence of the Morphological Range, Transition and Evolution of Stomatal Protection Mechanisms in Some Selected Proteaceae

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Evidence of the Morphological Range, Transition and Evolution of Stomatal Protection Mechanisms in Some Selected Proteaceae EVIDENCE OF THE MORPHOLOGICAL RANGE, TRANSITION AND EVOLUTION OF STOMATAL PROTECTION MECHANISMS IN SOME SELECTED PROTEACEAE Ratnawati Submitted in fulfilment of the requirements for the Masters of Science Degree l ' \ ' i. <.. I . t I \ I \ :'\. ' • SCHOOL OF PLANT SCIENCE DECEMBER 2001 DECEMBER 2001 This thesis is not to be made available for loan or copying for two years following the date this statement was signed. Following that time the thesis may be made available for loan and limited copying in accordance with the Copyright Act 1968. - - ---- - __·1 -----------~--- --- -- ------------- ------- DECLARATION Except as stated herein, this thesis contains no material which has been accepted for the award or any other degree or diploma, and to the best of my knowledge and belief contains no copy or paraphrase of material previously published or written by any other person, except where due reference is made in the text. I dedicate my work to my beloved husband, Agung, and my sons, Odit and Yusta, for their spiritual support during my study. Abstract Xero- and scleromorphic adaptations are obviously shown by Australian plants, in response to the Australian climate and edaphic factors. Since these adaptations overlap, there are problems separating the two. Some qualitative hypotheses about the distinction between xero- and scleromorphic characters have been proposed. This research is an effort to quantitatively determine xeromorphic characters in some members of the Proteaceae, in order to elaborate upon some of the existing hypotheses about these characters. Twenty three species of Banksia, 16 species of Grevillea and 6 species of Orites were sectioned and observed under the light micrscope and measurements were made of the stomata! depressions, margin recurvations, cuticle thickness and hair dimensions. Cuticle peels were made in order to count the number of stomates and hairs on the leaf surface. From the characters measured the following were calculated: the depression index (Id), recurved margin index (Irm), epidermis index (le), hair. index _(Ih) and cuticle components. The sum totals of the calculations were called the stomata! protection component (Spc). These data were then analysed using an analysis of variance with five replicates per species, in order to investigate whether there were any differences in characters determined .among species observed. Every index calculated (Id, Irm, Ih, le) and the cuticle components were analysed separately, with the aim of determining the intensity of the association between the characters observed and the minimum of the range of habitat annual rainfall, the maximum of the range ·of habitat annual temperature, the maximum of the range of habitat daily radiation and the maximum of the range of maximum daily wind run. An analysis of correlation was applied to these data. A similar method was applied for observations on juvenile and adult leaves. On the basis of these results the possible evolution of Banksia is reconstructed. Results showed that in Banksia species stomata! depression seems to have an important role protecting the stomates from the effect of habitat climatic factors, especially from temperature and radiation effects. The presence of recurved margins and hairs, however, were not as important as stomata! depressions. In Grevillea species, cuticle characters had a significant role in stomata! protection, particularly the stomata! and lower epidermis cuticle. A _contradictory finding to that of Banksia in this genus was that stomata! depression did not contribute to the stomata! depression. In comparison to the two genera already discussed, Orites was quite different. However, the hairs had a significant contribution in protecting stomates from temperature and radiation effects. - The phenomenon of leaf transition from juvenile to adult in some species observed showed some performance similarities to plants grown in harsh environments. Banksia blechnifolia leaves exhibited more intense stomata! depression and denser leaf hairs in adult leaves. It was assumed that these structures increased stomatal protection from excessive radiation. Similar growing phenomena were displayed by leaves of Grevillea pyramidalis. Stomata! depression in this species developed from being not present to being present and then deepened in later leaves. In addition, leaf hairs showed a gradual alteration in density from seedling through juvenile and adult leaves. However, leaves of B. marginata did not show the presence of stomata! depressions either in juvenile or adult stage. The leaf hairs, however, exhibited similar phenomena to those found in B. blechnifolia and G. pyramidalis. The results suggest that stomatal depression, margin recurvation, leaf hairs and cuticle characters increase stomata! protection either individually or collectively. It thus can be concluded that there is a possibility that these characters are xeromorphic when expressed by Banksia, Grevillea and Orites species . .. ll Acknowledgements First of all, I would like to deliver my special acknowledgement to my supervisor, Prof. Robert S. Hill, for his guidance, encouragement, and assistance with my writing, and with my finances at the end of my candidature. Another special acknowledgement goes to Dr. Anthony Koutoulis, for permitting me to work in his Tissue Culture Laboratory and lending me the electronic microtome and other equipment during my experiments. Other people who I would especially like to thank are: -My room mate Richard Barnes for giving me his hands and ideas since my early candidature. -Dr. Gregory Jordan, for his highly valued assistance in statistical analysis and BIOCLIM analysis. -My room mate Sutrisno for his extra assistance in computer operation and statistical analysis, and some brilliant ideas for my report. -Tang Yifeng for his extra help in operating the computer and as a field companion. -Nathalie Brown and Susan Lennon for their guidance in preparing resin embedding sections. -Lisa Schimanski for her patient help in working with ESEM. -Ian Cummings and Tracey Jackson for providing their warm hands for my plants in the glasshouses. -Katie Dutch, for helping me during my work in the Palaebotanical Laboratory. -Gaye Johnson for her assistance in organising my orders. -Dr. David Steele from the ESEM Laboratory, for his valuable assisstance in specimen gold coating and sending ESEM images. -Margaret Eldridge for proofreading my thesis. -Librarians in the Geography Department for their assistance during the tough time collecting locality data. -Staff of Nindethana Seed Service Pty, Ltd., Western Australia, for providing me with some Banksia and Grevillea seeds. - - -Neil Marriot from Stawell, Victoria, for providing me with some Grevillea foliage. -Deborah Edwards from the Australian National Botanical Gardens, Canberra, for sending me some Banksia and Orites foliage. -David Stewart from the Royal Botanic Gardens, Carnbourne, for providing me with some Banksia foliage. -Staff of The Parks and Wildlife Service, for giving me permits to get Orites seeds - - and foliage from Mount Field. Ill -Ms Jo Palm.er from The Centre for Plant Biodiversity for sending me locality data of some Proteaceae species I observed. -Paul Robin from The Centre for Botanical Research and Information on the Queensland Flora, for sending me locality data of some Proteaceae species I observed. -Last but not least, special thanks to Meity, Refli, members of the Indonesian Student Association, Diane Sward, Craig Gilmour, Ivonne Vucica, Dr. Robert C. Elliot, Dr. Tim Brodrib and Dr. Mark Hovenden for their warm friendship and help during my candidature. The very last special thanks I would like to give to my dearest husband and my boys, for their patience, caring and happiness during my three year study; to my flat mate Michael J. Freeland with his entertaining attitude and white coffee; and to my friend Sutrisno, for his company during the last months of my writing. ''" Table of contents Abstract .................................................................................................................... i Acknowledgements ............................................................................................... ii Table of contents ................................................................................................... iv Chapter 1. General introduction ............................. ..... .. .... ...... .......... ...... ... ... .. 1 Chapter 2. The morphological range of stomata! protection mechanisms in some selected Proteaceae .. .. .. ...... ... .. .. .. .. .. .. .. ... .. ... ... .. ... .. .. .. .. 7 2.1. Introduction ............................................................................. 8 2.2. Materials and methods .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 27 2.2.1. Sampling ........................................................................ 27 2.2.1.1. Banksia ................................................... :........ 27 2.2.1.2. Grevillea .......................................................... 31 2.2.1.3. Orites .............................................................. 34 2.2.2. Resin embedding section .. .. .. .. .. .. .. .... .. .. .. .. .. .. ... .. .. 35 2.2.3. Cuticle preparation for transmitted light microscopy . .. .. ... .. .. .. .. 36 2.2.4. SEM preparation for morphological observation .. .. .. ... .. .. .. .. .. .. .. .. .. .. .. .. .. .. 36 2.2.5. Data collection .............................................................. 37
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