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Starch Function in Ornamental Tobacco Floral Nectary Development Gang Ren Iowa State University

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Starch Function in Ornamental Tobacco Floral Nectary Development Gang Ren Iowa State University Iowa State University Capstones, Theses and Retrospective Theses and Dissertations Dissertations 2007 Starch function in ornamental tobacco floral nectary development Gang Ren Iowa State University Follow this and additional works at: https://lib.dr.iastate.edu/rtd Part of the Botany Commons, and the Plant Biology Commons Recommended Citation Ren, Gang, "Starch function in ornamental tobacco floral nectary development" (2007). Retrospective Theses and Dissertations. 15979. https://lib.dr.iastate.edu/rtd/15979 This Dissertation is brought to you for free and open access by the Iowa State University Capstones, Theses and Dissertations at Iowa State University Digital Repository. It has been accepted for inclusion in Retrospective Theses and Dissertations by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. Starch function in ornamental tobacco floral nectary development by Gang Ren A dissertation submitted to the graduate faculty in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Major: Genetics Program of Study Committee: Robert Thornburg, Major professor Martha James Basil Nikolau Drena Dobbs Kang Wang Iowa State University Ames, Iowa 2007 Copyright © Gang Ren, 2007. All rights reserved UMI Number: 3259503 UMI Microform 3259503 Copyright 2007 by ProQuest Information and Learning Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. ProQuest Information and Learning Company 300 North Zeeb Road P.O. Box 1346 Ann Arbor, MI 48106-1346 ii Table of contents List of figures xii List of tables x Acknowledgements xi Abstract xiii Chapter 1 General introduction 1 1.1 Hypothesis 3 1.1.1 Hypothesis 1 5 1.1.2 Hypothesis 2 5 1.2 Plant floral nectary and nectar 6 1.2.1 Plant nectar 7 1.2.2 Nectar secretion models 8 1.2.3 Starch in floral nectaries 10 1.3 Starch metabolism in higher plants 11 1.3.1 Starch biosynthesis 13 1.3.2 Starch degradation 20 1.4 Outline of this dissertation 21 Chapter 2 Starch accumulation in ornamental tobacco floral nectary 23 Abstract 25 2.1 Introduction 26 2.2 Materials and methods 27 2.2.1 Materials 27 2.2.2 Plants 27 2.2.3 Transport of radiolabeled sugars 27 2.2.4 Carbohydrates analysis 28 Thin layer chromatography of radiolabeled sugars 28 Nectar carbohydrate composition 28 iii 2.2.5 Starch analysis 29 Isolation of starch from nectaries 29 Starch quantitation 29 Starch structural analysis 29 Amylose content analysis 30 2.2.6 Light Microscopy 31 2.2.7 Scanning Electron Microscopy 32 2.3 Results 32 2.3.1 Nectary development 37 2.3.2 Starch in nectaries 40 2.3.3 Starch is dynamically processed during nectary development 45 2.3.4 Role of transported photosynthate in nectar production 51 2.3.5 Is tobacco a model for other plant species? 59 2.4 Discussion 59 2.4.1 Transient nature of the nectary starch 62 2.4.2 Role of transported photosynthate in nectar production 63 Chapter 3 Regulation of starch metabolism in ornamental tobacco floral nectary 69 Abstract 71 3.1 Introduction 72 3.2 Materials and methods 74 3.2.1 Plant Materials 74 3.2.2 rDNA methods 74 3.2.3 PCR methods 75 Primer design 75 General PCR 75 RT-PCR detection 75 Quantitative RT-PCR detection 76 3.2.4 Bioinformatics tools 77 3.2.5 Microscopy methods 77 iv Transmission Electron Microscopy 77 Quantitation of plastid volume 78 3.2.6 RNA isolation 78 3.2.7 protein methods 79 3.2.8 Real time RT-PCR detection 82 Protein isolation and enzyme activity assay 79 Protein isolation and immunodetection 79 3.3 Results 80 3.3.1 Starch metabolic genes 84 3.3.2 Regulation of starch biosynthesis 97 Sucrose synthase in developing ornamental tobacco floral nectary 97 ADP-glucose pyrophosphorylase in developing floral nectaries 99 3.3.3 Additional starch metabolic genes in developing floral nectaries 102 Starch synthases 102 Starch branching enzymes 105 Starch debranching enzymes 105 3.3.4 Quantitative RT-PCR (qRT-PCR expression) 108 3.4 Discussion 113 Chapter 4 β-carotene meatabolism in ornamental tobacco floral nectary 125 Abstract 127 4.1 Introduction 128 4.2 Materials and methods 129 4.2.1 Plant material 129 4.2.2 Light microscopy 129 4.2.3 Vibratome sections 130 4.2.4 Electron microscopy 130 4.2.5 Biochemical analyses 131 Carotenoids analysis 131 Ascorbic acid analysis 131 v 4.2.6 Labeling of carotenoids with [14C]-sucrose 132 4.2.7 Oligonucleotides for RT-PCR analysis 132 4.3 Results 133 4.3.1 Light Microscopy: S9 through S12 133 4.3.2 Electron microscopy: S9 through S12 138 S9 plastids 138 The S10 and S11 plastids 141 The S12 plastids 141 4.3.3 Biochemical and Molecular Genetic Analyses 144 4.4 Discussion 165 Chapter 5 Future work 170 5.1 Introduce of transgenic plants based future work 170 5.2 Materials and methods 179 5.2.1 Plante lines 179 5.2.2 RNAi constructs 179 5.2.3 Overexpression constructs 183 5.2.4 Plant transformation and plant selection 183 RNAi expression transgenic plants 183 Overexpression transgenic plants 183 Seeds collection 183 5.2.5 Genomic DNA verification of transgenic plants 184 Plant genomic DNA extraction 184 Genomic DNA PCR 184 5.2.6 Protein isolation 184 5.2.7 Western blot assay 185 5.2.8 Starch staining analysis 185 5.2.9 nectar carbohydrate analysis 185 5.3 Results 186 5.3.1 Transgenic plants verification 186 vi 5.3.2 Starch in ornamental flowers 186 Chapter 6 Summarization and discussion 198 6.1 Starch and floral development 200 6.2 Starch and nectar production 200 6.3 Starch biochemistry in developing tobacco floral nectary 202 Appendices 205 Appendix A Index of EST clones 206 Appendix B Index of oligonucleotides 211 Appendix C Index of constructs 220 Appendix D Index of transgenic plants 225 References 234 vii List of figures Figure 1.1 Structure of tobacco flowers 2 Figure 1.2 The core pathway of starch biosynthesis in higher plants 14 Figure 1.3 The core pathway of starch biodegradation in higher plants 15 Figure 1.4 Biochemical reaction of starch metabolic enzymes 16 Figure 2.1 Light and scanning electron microscopy (SEM) graphs of Arabidopsis thaliana and ornamental tobacco floral nectaries 34 Figure 2.2 Events occurring during development of tobacco nectaries 38 Figure 2.3 Light microscope images of 1-μm-thick LR White resin sections of tobacco nectaries at five different developmental stages stained for differential contrast enhancement with toluidine blue O or PAS technique for cell walls and starch 41 Figure 2.4 Accumulation of starch in nectary 43 Figure 2.5 FACE analysis of nectary starch 46 Figure 2.6 Amylose/amylopectin content of starch isolated from ornamental tobacco floral nectaries at various stages of development 48 Figure 2.7 TLC analysis of radiolabeled nectar 53 Figure 2.8 Labeling of C14-sucrose in various floral stages 55 Figure 2.9 Light microscopic images of 1-μm-thick LR White resin sections of Arabidopsis thaliana floral nectaries stained with PAS technique 60 Figure 2.10 Comparison of biochemistry with nectary development 66 Figure 3.1 Starch granule changes in developing floral nectary parenchyma cells 81 Figure 3.2 Neighbor-joining trees for ornamental tobacco starch metabolism genes 91 Figure 3.3 Sucrose synthases (SuSy) expression in ornamental tobacco 100 Figure 3.4 AGPase expressions in ornamental tobacco floral nectary 103 Figure 3.5 RT-PCR detection of starch metabolism related gene expression profile in ornamental tobacco 106 Figure 3.6 Real-time RT-PCR detection of starch metabolism related gene expression in ornamental tobacco floral nectaries 110 viii Figure 3.7 Model of starch mass accumulation and starch metabolic enzyme activity in ornamental tobacco floral nectary 119 Figure S3.1 Comparison of splice-variant of α–amylase 121 Figures 4.1-4.18 Images of isolated whole, and 30-µm-thick cross and longitudinal vibratome sections of tobacco gynoecia and basal nectaries from Stage 2 through Stage 12 observed with normal light or between crossed polarizers 136 Figures 4.19-4.24 Light and transmission electron microscope sections through nectary at Stages 9-12 139 Figures 4.25-4.31 Transmission electron microscope sections showing nectary cell plastids from S9 through S12 142 Figure 4.32 Identification of ornamental tobacco nectary carotenoid as β-carotene 147 Figure 4.33 Analysis of nectary expressed carotenoids extracted from LxS8 149 Figure 4.34.RT-PCR of gene encoding β–carotene biosynthetic enzymes 153 Figure 4.35 Interconnection of biochemical pathways in the nectary 156 Figure 4.36 Thin layer chromatogram (TLC) visualization of nectary expressed carotenoids 159 Figure 4.37 Quantity of ascorbate extracted from six nectaries of ornamental tobacco flowers at different developmental stages 161 Figure 4.38 Ascorbate metabolism 163 Figure 5.1 Scheme of transgenic plants 171 Figure 5.2 3D structure of the DNA-binding domain of sucrose responsive element binding proteins 175 Figure 5.3 Gene expression pattern of SREBP detected by RT-PCR 177 Figure 5.4 Gene expression of mevalonate pathway and MEP pathway genes detected by RTPCR 181 Figure 5.5 Genomic DNA PCR verification of transgenic plants 187 Figure 5.6 Immunodetection of transgenic plants 189 Figure 5.7 Starch accumulation detected by iodine staining in LxS8 flowers 191 ix Figure 5.8 Starch staining of various transgenic plants line in ornamental tobacco 194 Figure 5.9 Nectar carbohydrates measurement of various transgenic plants line in ornamental tobacco 196 Figure 6.1 Speculated biochemical process in ornamental tobacco floral nectary development 203
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