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The Effect of Paclobutrazol on Flowering Activity and Gibberellin

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The Effect of Paclobutrazol on Flowering Activity and Gibberellin The effect of paclobutrazol on flowering activity and gibberellin levels in Eucalyptus nitens and Eucalyptus globulus by Omar Hasan B.Sc. Submitted in fulfilment of the requirements for the degree of Doctor of Philosophy University of Tasmania, Hobart October, 1993 Declaration This thesis contains no material that has been accepted for the award of any other degree or diploma in any university and contains no copy or paraphrase of material previously published or written by another person, except where due reference is made in the text. OmarHasan 12th of October, 1993 ii For my grandmother, A.E. Dawson. ill Abstract Experiments prior to this project demonstrated the capacity of the plant growth retardant paclobutrazol to enhance flowering in commercially important Eucalyptus nitens and E. globulus trees and produced anecdotal evidence of this material reducing time to first flowering. Paclobutrazol is known to reduce the levels of endogenous gibberellins (GAs) in several species and hence it was hypothesized that the effects of paclobutrazol on flowering in these Eucalyptus species may be mediated by an effect on GA levels. The lack of previous identification of GAs in this genus necessitated the development of extraction and purification procedures to identify and quantify GAs. The compounds identified suggested that the early 13- hydroxylation pathway was the dominant mechanism for production of GAs, with GA 1 as the likely biologically active compound. Persistence of a paclobutrazol induced increase in flowering of grafted E. nitens material was related to the continued depression of endogenous GA concentrations. Additionally, higher concentrations of GA1 were found to be correlated with reduced flowering responses in this reproductively competent material. A lowering of endogenous GA levels, in combination with a co-requisite of a period of cold has been associated with the induction of first flowering in newly grafted E. nitens material. The effects of cold treatment were not mediated by an effect on levels of GA1 or GA2o· Soil applied paclobutrazol was shown to travel up stems and accumulate in leaf tissue. Breakdown in plant tissue was shown to be rapid, with a half-life likely to be substantially less than 21 d. Soil and foliar application methods were shown to produce different patterns of metabolism of labelled paclobutrazol, as demonstrated by HPLC separation of labelled metabolites extracted from growing apices. The rate of breakdown in the soil was observed to be variable, but slow in comparison to that within plant tissues and may be the source of the considerable persistence of effects of paclobutrazol application observed in some field trials. Application of paclobutrazol to 6 month old E. globulus seedlings resulted in the production of flower buds at less than 18 months of age despite the retention of juvenile foliage. One year later, following normal bud development, anthesis and pollination, capsules were produced, while maintenance of material in a range of iv growth conditions over the second winter again demonstrated the strong requirement for cold seen in grafted E. nitens, as well as revealing an apparent promotion of flowering associated with reduced pot size. The reduction in generation time achieved using commercial seedlings was ca. 50% (3 years) which should be of major benefit to tree breeders, given that the long generation time of eucalypts is a major determinant of the rate at which genetic gains can be made by conventional tree breeding methods. In reproductively competent seedlings and grafts, paclobutrazol application was confirmed to increase the average number of flower buds per seedling. This could be advantageous when seed requirements from an elite tree are high, or when yearly seed yield tends to be variable. v Acknowledgements I would like to thank the following people for the help they have provided over the course of my post-graduate studies. -Prof. Jim Reid for supervision and guidance through all aspects of post­ graduate training. -Dr John Ross for his plentiful assistance and the willingness with which it was given. - Mr Mike Moncur of CSIRO Division of Forestry, Canberra for his cheerful help and his tireless work in our fruitful (sic) collaboration (see Chapter 2). - Sandra Hetherington, formerly of Forest Resources and now with Australian Newsprint Mills, for providing me with plant material on very short notice on more than one occasion and for keeping me updated on the progress of her fme work. -Peter Naughton of Forest Resources and David deLittle, Ian Ravenwood and Wayne Tibbits of Associated Pulp and Paper Mills for their friendly help and interest. - Mr Noel Davies for his completely invaluable assistance with mass­ spectrometry of gibberellins. -Dr Brad Potts for providing eucalypt seed and helping with experimental design and statistical analyses. -Dr Gregory Jordan for help with statistical analyses. -Drs Robert Wiltshire and Dick Pharis for various helpful suggestions. -Leigh Johnson, Peter Bobbi and Kathy McPherson in Hobart and John Turner in Canberra for maintaining experimental material. - Heidi Dungey and Jon Marsden-Smedley for tolerating me as a room mate. vi &knowledgements - Stephen Swain, Christine Beveridge, Jim Weller, Mike Battaglia, Kristen Williams and Naomi Lawrence for their help and company as fellow Ph.D. students. - Peter Gore, Andrea Manson, Rhyl Opie, Paula Chalmers, Kate Booth and Dale Barron for help provided as part of their 3rd year Plant Science curriculum. - Erik Wapstra for his competent help in preparation of this thesis. - The Plant Science Tea Room cribbage players for many hours of entertainment. - My parents for their support and encouragement throughout my years of schooling. - And finally Lisa Stanton for her companionship and understanding. V1l Table of Contents Title ~huation ii Dedication iii Abstract iv Acknowledgements vi Generallntroduction 1 What is a Plant Growth Retardant 2 Varieties of Plant Growth Retardants 3 Effects of Plant Growth Retardants on GA Biosynthesis 5 Effect of Plant Growth Retardants on other Biosynthetic Pathways 6 Plant Growth Retardant Effects 8 Optimisation of Plant Growth Retardant Effects in Woody Angiosperms 11 Plant Growth Retardants in Phytohormone Research 13 Conclusion 14 Aims of the Present Study 14 Figures 17 Chapter 1- Identification and Quantification of GAs in E. nitens 19 Tables 32 Figures 34 Chapter 2 - Effects of Paclobutrazol and Cold on GA Levels and Flowering in Grafted E. nitens 40 Tables 50 Figures 55 Plates 58 Chapter 3 - Paclobutrazol Movement and Metabolism 62 Figures 78 Chapter 4 -Induction of Flowering Precocity in E. globulus Seedlings 88 Tables 105 Figures 110 Plates 119 General Discussion 123 Figures 134 Plates 135 References 142 Appendix A- Hasan et al. (1994) 163 Appendix B -Methods section of Moncur et al. (1993) 183 Appendix C - List of published works 186 viii 1 General Introduction Eucalyptus globulus (Labill.) and Eucalyptus nitens (Deane and Maid.) Maid. are commercially important species of the myrtaceous genus Eucalyptus. They are heavily utilised for the production of wood pulp both in Australia (eg. Tibbits, 1986) and overseas (Zacharin, 1978) and are the most widely planted temperate species of eucalypts, which is in tum the most widely planted genus of angiosperm trees (Griffin, 1989). To obtain maximum gain from such a scale of planting requires large quantities of seed, preferably from improved trees. Work using plant growth retardants on orchard species such as apples (Tukey, 1981; Jones et al., 1989a) and citrus species (Mauk et al., 1986) was observed to increase bud numbers in the seasons following application, leading to an interest in assessing their effects on the reproductive output of eucalypt species. If fecundity of desirable trees could be increased within a seed orchard, a larger amount of improved seed would be available to satisfy the requirements of new/renewed plantations. It was also of interest to assess the capacity of plant growth retardants to affect the relatively long generation times notable for species of this genus (Ashton, 1975; Turnbull and Pryor, 1978). These long generation times pose an important problem to eucalypt breeders (Griffin, 1989) due to the limitations on rates of incorporation of new traits they impose. A capacity for precocious (early) flowering under natural conditions has been noted for several species of eucalypts (Pryor, 1966). If a chemical or cultural treatment could be established to promote the expression of this capacity, breeding programs would benefit greatly, as long as the reproductive development of the treated plant continues to occur normally. In conifers, there are many reports of successful promotion of both heavier and precocious flowering through the use of cultural techniques ranging from stem girdling, to release from competition by stand thinning (Owens and Blake, 1985) and/or chemical treatments by application of growth retardants, especially non­ polar gibberellin (GA) mixtures such as GA417 (Pharis and King, 1985; Pharis et al., 1987). Such a level of control over flowering processes in eucalypts and other woody angiosperms is the ultimate practical goal sought by tree breeders, while gaining an understanding of the control of flowering is a fundamental aim Qeneral I ntrQd!JCtion 2 from a theoretical standpoint. As work to this stage has suggested that the use of plant growth retardants may be an effective means of promoting flowering in woody angiosperms including eucalypts, an
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