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Photosynthetic Adjustments of Amaranthus Varieties In PHOTOSYNTHETIC ADJUSTMENTS OF AMARANTHUS VARIETIES IN RESPONSE TO GROWTH IRRADIANCE A Thesis Submitted to the College of Graduate and Postdoctoral Studies In Partial Fulfillment of the Requirements For the Degree of Master of Science In the Department of Plant Sciences University of Saskatchewan Saskatoon By Elena Benic © Copyright Elena Benic, August 2018. All rights reserved. PERMISSION TO USE In presenting this thesis in partial fulfillment of the requirements for a Postgraduate degree from the University of Saskatchewan, I agree that the Libraries of this University may make it freely available for inspection. I further agree that permission for copying of this thesis in any manner, in whole or in part, for scholarly purposes may be granted by the professor or professors who supervised my thesis work or, in their absence, by the Head of the Department or the Dean of the College in which my thesis work was done. It is understood that any copying or publication on use of this thesis or parts thereof for financial gain shall be given to me and to the University of Saskatchewan in any scholarly use which may be made of any material in my thesis. Requests for permission to copy or to make other use of materials in this thesis in whole or in part should be addressed to: Head of the Department of Plant Sciences University of Saskatchewan Saskatoon, Saskatchewan S7N 5A8 Canada OR Dean College of Graduate and Postdoctoral Studies University of Saskatchewan 107 Admission Place Saskatoon, Saskatchewan S7N 5A2 Canada i ABSTRACT Increased concern of food security has created a demand for crops that exhibit high yields under conditions that require a minimal input of resources, such as light. Amaranthus is a C4 plant that has multiple uses, including that of a food source in many parts of the world. The objective of this research was to evaluate the photosynthetic performance of red and green vegetable varieties of Amaranthus blitum grown under high light (HL; 500 µmol m-2 s-1) photosynthetic photon flux density (PPFD) and low light (LL; 70 µmol m-2 s-1 PPFD). Under HL growth irradiance, the red variety accumulated more biomass than the green variety before flowering. This was accompanied by a 1.2- fold increase of photosynthetic efficiency, a 1.5-fold increase of photosynthetic capacity (maximal rates of O2 evolution), a 1.5-fold increase of dark respiration rate, a 1.3- fold increase in light compensation point and a 1.7-fold increase of light saturation point. These values were supported by a 2.5- and 2.6-fold higher content of chlorophyll and carotenoids, respectively. A 4.7-fold greater betalain content was also observed. In addition, exposure to a photoinhibitory irradiance (1450 µmol m-2 s-1 PPFD) at 2°C for 4 hours demonstrated the red variety exhibited increased tolerance to photoinhibition in comparison to the green variety when measured at the level of maximal photochemical efficiency of photosystem II. Further, this may be a result of the screening of light and shielding of chloroplasts by betalains that accumulate predominantly in the abaxial mesophyll and supported by a 1.2-fold increase in the red variety lower mesophyll cells thickness than in the green variety, leading to a thicker leaf. The granal index in bundle sheath cells under high light irradiance in the red variety was 2.0-fold greater than in the green variety, supported by 3.1-fold greater ratio of the length of appressed/non- appressed thylakoids and a 2.0-fold increase in thylakoids/granum. These results suggest that under HL growth irradiance, the granal index and stackness (thylakoids/granum) enhances photosynthetic efficiency and capacity. In contrast, under LL growth irradiance both varieties performed photosynthetically similarly. Collectively, these results indicate that the red variety possesses a greater photoacclimatory capacity than the green variety under HL growth irradiance, whereas neither variety expressed an advantage under the LL growth irradiance in this study. ii ACKNOWLEDGEMENTS I would like to express my deepest gratitude and appreciation to Dr. Gordon R. Gray and Dr. Karen K. Tanino for their encouragement, scientific guidance, patience and financial support during my studies. I would also like to thank my graduate committee members: Dr. Rosalind A. Bueckert (chair), Dr. Robert H. Bors and Dr. Art Davis for providing their diverse backgrounds to support me during this project, as well as deep thanks to Dr. Simon D. X. Chuong for being the External Examiner and for his imput to the completion of this work. Special thanks to the staff of the Department of Plant Sciences, especially: Dr. Yuguang Bai (head), Carolyn Ouellet, Gloria Gingera and Marlene Freeman. Special thanks to the many members of Dr. Tanino’s Lab for their time, advices and support with specific note of Dr. Ian Willick, Kaila Hamilton, Eric Rae, Dr. Prakash Venglat, Pankaj Banik, Gowribai Valsala, Rensong Liu, Dr. Masoomeh Etehadnia, Dr. Tawhidur Rahman and Abdul Halim. Quality plant care was made possible by the phytotron team. Special thanks to Dr. Guosheng Liu from Department of Biology for advice and help in preparation of samples for TEM, to Larhonda Sobchishin from WCVM imaging center using TEM. Microscopy techniques were learned from the imaging team at NMBU with special recognition of Drs. YeonKyeong Lee, Hilde Kolstad, Lene Cecilie Hermansen and Jorunn Olsen. Special thanks also to Dr. Chris Ambrose for his advice and support. Without the numerous and generous funding sources this project would have not been possible: Special thanks to Saskatchewan Ministry of Agriculture, Natural Sciences and Engineering Research Council of Canada, Saskatchewan Innovation & Opportunity Scholarship, Rene Vandeveld Postgraduate Scholarship and Daniel Geddes Graduate Scholarship in Plant Sciences, as well as the Department of Plant Sciences. Finally, I would like to recognize my husband Kresimir Benic, and friends (especially Tatiana Mashkova) for their interest and unwavering support over the course of the project. iii DEDICATION This thesis is dedicated to my beloved husband Kresimir Benic, son Jakov, and daughters Azra and Judita for their love, support and incredible patience. iv TABLE OF CONTENTS Page PERMISSION TO USE ............................................................................................... i ABSTRACT ................................................................................................................. ii ACKNOWLEDGEMENTS ........................................................................................... iii DEDICATION .............................................................................................................. iv TABLE OF CONTENTS .............................................................................................. v LIST OF TABLES ........................................................................................................ viii LIST OF FIGURES ...................................................................................................... x LIST OF ABBREVIATIONS ......................................................................................... xii 1.0 INTRODUCTION ................................................................................................... 1 1.1 Hypothesis and Objectives ......................................................................... 3 1.1.1 Hypothesis .................................................................................... 3 1.1.2 Overall Objective ........................................................................... 3 1.1.3 Specific Objectives ........................................................................ 4 2.0 LITERATURE REVIEW ......................................................................................... 5 2.1 Photosynthesis ........................................................................................... 5 2.1.1 Leaf Structure and Anatomy .......................................................... 5 2.1.2 Stomata and Trichomes ................................................................ 6 2.1.3 Chloroplasts .................................................................................. 8 2.1.3.1 Peripheral Reticulum ......................................................... 8 2.1.3.2 Cytoplasmic Protrusions .................................................... 10 2.1.3.3 Crystalline Inclusions ......................................................... 10 2.1.4 Microbodies ................................................................................... 10 2.1.5 Light Dependent Reactions ........................................................... 11 2.1.6 Photosynthetic Measurements ...................................................... 13 2.1.6.1 Photosynthetic O2 Evolution .............................................. 14 2.1.6.2 Chlorophyll Fluorescence .................................................. 14 2.1.7 Photosynthetic Carbon Reduction ................................................. 14 2.1.8 Photorespiration ............................................................................ 20 2.2 Photostasis and Photoinhibition .................................................................. 21 2.2.1 Photostasis .................................................................................... 21 2.2.2 Photoinhibition ............................................................................... 22 2.3 Photoacclimation .......................................................................................
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