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Carotenoid Biosynthesis and Metabolism in Potato Tubers

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Carotenoid Biosynthesis and Metabolism in Potato Tubers Carotenoid biosynthesis and metabolism in potato tubers Adriana Stefania Pasare A Thesis Submitted for the Degree of Doctor of Philosophy Royal Holloway College, University of London November 2012 Declaration of Authorship I hereby declare that the present thesis is of my own composition and based on the results of work conducted by myself. The results reported here have not, in whole or in part, been previously presented for a higher degree or qualification. For investigations other than my own, there is a clear indication in the text by reference to the relevant workers or their publications. Adriana Stefania Pasare ii Acknowledgements I would like to start by quoting Marie Curie, who said ‘I am among those who think that science has great beauty. A scientist in his laboratory is not only a technician: he is also a child placed before natural phenomena which impress him like a fairy tale’. It is a pleasure to thank all of those who made this thesis possible, thus giving me the opportunity to scientifically explore cellular processes and be humbly impressed with the correlations established by nature. I am heartily thankful to my supervisors, Mark Taylor, Alison Roberts, Paul Fraser and Peter Bramley, whose encouragement, guidance and support from the initial to the final level enabled me to develop a subject I am passionate about. Also, I would like to thank Kath Wright, Ray Campbell, Wayne Morris and Sean Chapman, for their constant support and helpful technical advice, but also all other JHI collaborating staff. Lastly, I can but offer my love and blessings to all my family, who have always believed in me, encouraging me through my years of study. Thank you, Mami, Oana, George, Johnny and my dear loving husband, Ionut. iii Abstract Potato tuber flesh colour is a quality trait dependent on the types and levels of carotenoids. These, together with many plant growth regulators, sterols and terpenes, are isoprenoid derived compounds. Even though carotenoids have been extensively studied due to their importance in photosynthesis and nutrition, a clear understanding concerning the regulation of their accumulation and localisation in plant storage organs is lacking. The present study describes two strategies to investigate the regulation of carotenoid biosynthesis in potato. Firstly, in order to localise the site of synthesis of tuber carotenoids, stable transgenic potato plants were generated. These expressed two key carotenoid biosynthetic genes, PHYTOENE SYNTHASE 2 (PSY2) and β-CAROTENE HYDROXYLASE 2 (CrtRb2), fused to red fluorescent protein (RFP). The expression and carotenoid levels were significantly increased, confirming functionality of the protein-RFP ensembles. Confocal microscopy on developing tuber tissue made it possible to obtain images of PSY2-RFP and CrtRb2- RFP localisation into amyloplasts. Secondly, the effect of carotenoid degradation was addressed as a potential regulator of accumulation, based on the recent discovery that a CAROTENOID CLEAVAGE DIOXYGENASE 4 (CCD4) reduces carotenoid levels in potato tubers. CCD8 is another important member of the CCD gene family, with a role in strigolactone biosynthesis. In this study, CCD8-RNAi knock out lines were characterised. Although CCD8 down-regulation did not significantly affect carotenoid content, a range of interesting above and below ground phenotypes was observed. Importantly, the study showed that the CCD8 gene product impacts on stolon branching, tuber development and tuber dormancy. iv Table of contents Declaration of Authorship ............................................................................................ ii Acknowledgements .................................................................................................... iii Abstract ....................................................................................................................... iv Table of contents .......................................................................................................... v List of figures .............................................................................................................. xi List of tables .............................................................................................................. xiv List of abbreviations ................................................................................................... xv Chapter 1 Introduction .............................................................................................. 1 1.1 Potato and characteristics ............................................................................. 2 1.1.1 Origins and diversity ................................................................................ 2 1.1.1 Morphology and growth characteristics of potato plants ......................... 4 1.1.2 Production and economic importance ...................................................... 4 1.1.3 Consumption and nutritional value .......................................................... 7 1.2 Carotenoids .................................................................................................. 9 1.2.1 Chemical structure and types of carotenoids ........................................... 9 1.2.2 Importance of carotenoids ...................................................................... 12 1.2.3 Carotenoids in potato tubers-natural variation and content ................... 16 1.3 Regulation of carotenoid content in storage organs ................................... 17 1.3.1 Biosynthesis of carotenoids in plants ..................................................... 18 1.3.2 Manipulation of carotenoid content ....................................................... 25 1.3.3 Carotenoid storage in plants ................................................................... 27 1.3.4 Degradation of carotenoids .................................................................... 34 1.4 Aims of the present study ........................................................................... 37 Chapter 2 Materials and Methods ........................................................................... 39 2.1 Plasmids ..................................................................................................... 40 2.1.2 pGENTR1a ............................................................................................. 40 v 2.1.3 pGEM-T-Easy ........................................................................................ 40 2.1.4 pK7FWG2 .............................................................................................. 42 2.1.5 pK7RWG2 ............................................................................................. 44 2.1.6 pP19K ..................................................................................................... 45 2.1.7 pGreen mGFP 5ER 3’OCS .................................................................... 45 2.1.8 pHELLSGate 8 ....................................................................................... 45 2.2 Bacterial strains .......................................................................................... 46 2.2.1 E. coli DH5α .......................................................................................... 46 ® 2.2.2 OneShot TOP10 Electrocomp™ E. coli .............................................. 46 2.2.3 ElectroMAX™ DH10B™ ...................................................................... 46 2.2.4 ElectroMAX™ Agrobacterium tumefaciens LBA4404 ......................... 46 2.2.5 AGL1/pBBR1MCS1_ Vir GNS4D ........................................................... 47 2.2.6 AGL1 pSOUP ........................................................................................ 47 2.2.7 DB3.1 E. coli .......................................................................................... 47 2.2.8 Preservation of bacterial cultures ........................................................... 47 2.3 Transformation of competent cells............................................................. 48 2.3.1 Transformation of competent E. coli by heat shock .............................. 48 2.3.2 Transformation of competent E.coli and Agrobacterium by electroporation.................................................................................................... 48 2.4 Culture media and antibiotics..................................................................... 49 2.4.1 Bacterial culture media .......................................................................... 49 2.4.2 Tissue culture media .............................................................................. 49 2.4.3 Buffers .................................................................................................... 50 2.4.4 Antibiotics .............................................................................................. 53 2.4.5 Hormones ............................................................................................... 54 2.4.6 Fluorescent dye staining ......................................................................... 55 2.5 Cloning systems ......................................................................................... 56 vi 2.5.1 T/A cloning system ................................................................................ 56 2.5.2 pGENTR1a cloning ................................................................................ 56 2.5.3 Gateway cloning system ........................................................................ 57 2.6 Plant
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