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Elucidating TOR Function in Kalanchoe Daigremontiana Plantlet

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Elucidating TOR Function in Kalanchoe Daigremontiana Plantlet Elucidating TOR Function in Kalanchoë daigremontiana Plantlet Formation A thesis submitted to the University of Manchester for the degree of Master of Philosophy in the Faculty of Biology, Medicine and Health 2019 Kirsty McCready School of Biological Sciences TABLE OF CONTENTS LIST OF FIGURES AND TABLES .................................................................................................. 4 LIST OF ABBREVIATIONS .......................................................................................................... 5 ABSTRACT ................................................................................................................................ 6 DECLARATION.......................................................................................................................... 7 COPYRIGHT STATEMENT ......................................................................................................... 8 ACKNOWLEDGMENTS ............................................................................................................. 9 1 Introduction ....................................................................................................................10 1.1 Plant growth and development in response to nutrient availability .............................10 1.2 The Plant TOR kinase complex ......................................................................................11 1.2.1 The Arabidopsis TOR gene ......................................................................................11 1.2.2 The TOR protein......................................................................................................11 1.2.3 The TOR complex ....................................................................................................12 1.2.4 REGULATORY ASSOCIATED PROTEIN OF AtTOR .......................................................13 1.2.5 LETHAL WITH SEC THIRTEEN 8 ................................................................................14 1.3 TOR inhibition in plants ................................................................................................14 1.3.1 Genetic alteration of TOR expression ......................................................................14 1.3.2 Chemical inhibition of TOR expression ....................................................................15 1.4 TOR function during plant development .......................................................................15 1.4.1 Embryogenesis .......................................................................................................15 1.4.2 Germination and Seedling Growth ..........................................................................16 1.4.3 Meristem development ..........................................................................................19 1.4.4 Plant and Leaf size ..................................................................................................21 1.4.5 Flowering ................................................................................................................21 1.5 Triggered pluripotency in the species Kalanchoë daigremontiana ...............................23 1.5.1 The Kalanchoë genus ..............................................................................................23 1.5.2 Plantlet formation in the species Kalanchoë daigremontiana ..................................24 1.5.3 K. daigremontiana plantlet formation shares features of organogenesis and embryogenesis ................................................................................................................25 1.5.4 Plantlet initiation requires organogenesis meristem maintenance genes ................25 1.5.5 Plantlet formation follows an embryo-like developmental program ........................26 1.5.6 Plantlet formation and the TOR kinase ....................................................................28 1.6 MPhil Aims....................................................................................................................30 2 Materials and Methods ..................................................................................................31 2 2.1 Plant Growth Conditions ...............................................................................................31 2.2 Treatment Conditions ...................................................................................................31 2.3 Cloning and Plant Transformation ................................................................................31 2.3.1 Primer Design .........................................................................................................31 2.3.2 DNA Extraction .......................................................................................................32 2.3.3 Vector Construction ................................................................................................32 2.3.4 Golden Gate Assembly ............................................................................................33 2.3.5 Kalanchoë daigremontiana Transformation ............................................................34 2.4 Genotyping ...................................................................................................................35 2.5 RNA Extraction ..............................................................................................................35 2.6 cDNA Synthesis, RT-PCR and qRT-PCR...........................................................................36 2.7 ß-Glucoronidase (GUS) Staining ....................................................................................37 2.8 Phylogenetic Tree Construction ....................................................................................37 2.9 Image acquisition and Data Analysis .............................................................................38 3 Results ............................................................................................................................39 3.1 TOR, RAPTOR and LST8 phylogenies are congruent with the species trees ...................39 3.2 KdTOR expression varied across plantlet developmental stages ..................................40 3.2 TOR::GUS expression mirrors qPCR pattern across developmental stages ...................43 3.2.1 promoterTOR::GUS lines were generated ................................................................43 3.3 Torin2 application to the leaf margin inhibited plantlet formation ..............................46 3.4 Genetic suppression of TOR abolished plantlet formation............................................47 3.4.1 35S::TORantisense construct was assembled ..........................................................47 ........................................................................................................................................48 3.4.2 35S::KdTORantisense transgenic lines were produced .............................................49 3.4.3 Repression of tor severely restricted plantlet formation .........................................49 3.4.4 35S::KdTORantisense displayed changes in phyllotaxy and meristem growth ..........53 3.5 35S::KdSnRK1antisense construct was assembled ........................................................54 4 Discussion .......................................................................................................................55 5 Conclusion ......................................................................................................................63 6 References ......................................................................................................................65 Final Word Count: 16480 3 LIST OF FIGURES AND TABLES Figure 1: The plant TOR Complex (TORC)……………………………………………………….…………..13 Figure 2: Upstream and downstream targets of the TOR Complex (TORC) in plants…..18 Figure 3: Phylogeny of the Kalanchoë genus………………………………………………………………24 Table 1: Golden Gate Modules amplified by PCR and ligated into pGEM®T-Easy……….34 Table 2: Primer pairs designed for qRT-PCR…………………………………………………..…………..36 Figure 4: Phylogenetic trees of the plant TORC1 components, constructed from amino acid sequence alignments of the a) TOR, b) RAPTOR1 and c) LST8 genes.…………………………………..40 Figure 5: Stages of Kalanchoë daigremontiana plantlet formation and expression of KdTOR and KdRAPTOR1……………………………………………………………………………….…………….42 Figure 6: Cloning of the KdTORpromoter region ~1.5 kb upstream of KdTOR and Agrobacterium tumefaciens transformation……………………………………………………..……….43 Figure 7: KdpTORpromoter::GUS expression during plantlet development…………....….45 Figure 8: Torin2 and mock treatment of Kalanchoë daigremontiana plantlet formation…………………………………………………………………………………………………………………..46 Figure 9: Generation of 35S::KdTORantisense…………………………………………………………….48 Figure 10: Genotypic and phenotypic analysis of 35S::KdTORantisense………………..……50 Figure 11: Microscope images of 35S::KdTORantisense plantlet development stages..52 Figure 12: Phyllotaxy and meristem phenotypes in 35S::KdTORantisense………………….53 Figure 13: Generation of 35S::KdSnRK1antisense……………………………………………………….54 4 LIST OF ABBREVIATIONS 5’TOP 5’ Terminal Oligopyrimidine tract 5’ uORF 5’ upstream Open Reading Frame ABA Abscisic Acid ABI3 ABSCISIC ACID INSENSITIVE 3 AP1/2/3 APETALA 1/2/3 ARF AUXIN RESPONSE FACTOR asTORi active site TOR inhibitor bHLH basic Helix-Loop-Helix BIN2 BRASSINOSTEROID INSENSITVE 2 BPEp BIGPETALp bZIP11 Bovine Serum Albumen CAM Crassulacean Acid Metabolism
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