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Conifers Response to Water Stress: Physiological Responses and Effects on Nutrient Use Physiology

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Conifers Response to Water Stress: Physiological Responses and Effects on Nutrient Use Physiology CONIFERS RESPONSE TO WATER STRESS: PHYSIOLOGICAL RESPONSES AND EFFECTS ON NUTRIENT USE PHYSIOLOGY By İsmail Koç A DISSERTATION Submitted to Michigan State University in partial fulfilment of the requirements for the degree of Forestry – Doctor of Philosophy 2019 ABSTRACT CONIFERS RESPONSE TO WATER STRESS: PHYSIOLOGICAL RESPONSES AND EFFECTS ON NUTRIENT USE PHYSIOLOGY By İsmail Koç Conifer species are the most extensively distributed on earth, and they are one of the most significant renewable resources with high economic value. Conifer species, Pinus and Abies species have been gaining popularity due to their desirable green color for products such as Christmas trees and are extensively used in landscaping. Not only inhabiting forest in their natural habitat, but also in plantations and reforestation areas usually outside their natural range where they have been exposed to water stress due to water shortage and the effects of climate change. Water stress is an important environmental factor for tree growth and development in plants. Therefore, we investigated the effect of irrigation and fertilization on balsam (Abies balsamae) and concolor fir (Abies concolor) and white pine (Pinus strobus) seedlings in terms of tree morphology and physiology using a factorial design with three species and irrigation levels and two fertilization rates. Increased irrigation not only increased morphological traits such as diameter and height growth but also increased the net photosynthesis and stomatal conductance. The combination of each treatments had 5 seedlings for fir species and 4 seedlings for the pine species totaling 168 individual trees. White pine and balsam fir showed some drought tolerant mechanisms where concolor fir exhibited drought avoidance mechanisms. Fir species had higher net photosynthesis, stomatal conductance, transpiration rate and a lower water use efficiency compared to white pine. White pine had lower potassium concentration compared to two fir species, and balsam fir had higher calcium concentration compared to white pine and concolor fir under stress conditions, implying that fir species are more susceptible to water stress. We observed that concolor fir had a greater capacity for conserving water compared to white pine, leading to better above ground growth and shoot to root ratio. Balsam and concolor fir also had a greater foliar nitrogen concentration compared to white pine seedlings due to using an avoidance mechanism and maintaining nutrient uptake under water deficit conditions. White pine trees use drought tolerance strategies to reduce transpiration and maximize water uptake with increased root systems. White pine trees had higher below-ground biomass, with increased fine and coarse roots, and a lower nutrient use efficiency compared to two fir species. Moreover, we also tested the provenance and altitudinal variation of Turkish fir seedlings under water stress conditions. Morphological traits, such as relative root collar diameter, relative height growth, and stem volume index differed with seed source altitude as transplants from higher seed sources altitudes had greater growth compared to seedlings from lower altitudes. Overall, provenance had little effect on physiological parameters such as net photosynthesis, stomatal conductance, transpiration rate, internal CO2/ambient CO2 ratio, water use efficiency (WUE=A/E) and intrinsic water use efficiency (WUEi = A/gs), chlorophyll fluorescence (Fv/Fm) and carbon isotope discrimination rate (Δ13C). Provenances varied in stem water potential and net photosynthesis. Seedlings from the Karabuk provenances had high stem water potential and net photosynthesis. Intrinsic water use increased with altitude of the seed sources as seedlings from higher altitudes showed higher values compared to lower altitudes. Karabuk provenances might be more sensitive to water availability than Adapazari provenances. Adapazari provenances should be selected for the plantation and afforestation areas and production of Christmas trees. Copyright by İSMAİL KOÇ 2019 Dedicated to my grandfathers, Ekrem Koç and Ahmetcan Atmaca. You both were the first to teach me a lot about plants and their practices. I miss you, both. vi ACKNOWLEDGEMENTS I would first and foremost like to thank my major professor, Dr. Pascal Nzokou for giving me the opportunity to work with him on this project. He did a wonderful job not only supporting and encouraging me, but also providing me with the knowledge, guidance, and support. This project has been a very good learning experience and has helped me expand my knowledge and thought process immensely. I am also thankful for committee members Dr. Bert Cregg, Dr. Sophan Chhin and Dr. Paolo Sabbatini who spent many hours reading my dissertation and also for providing guidance and help during my project. I would like to thank the farm managers Randy Klevickas and Paul Bloese at the Tree Research Center for their patience and assistance. I am thankful for the help of my fellow colleagues Patrick Shults, Chad Papa, Dilek Yildiz, and several undergraduate assistants I worked with through the course of this project. I am sincerely grateful for the friendship and emotional support given by Salih Armagan, Cem Celal Tutum, Baburhan Uzum, Alp Ozdemir, Xinyue Cui, Nihan Çetin, Murat Uzel, Beyaz Basak Koca, Acimovic family, and all of my volleyball and soccer friends. Most important I would like to thank you my mother, father, and parents for supporting my decision to continue my education and the support they provided for me during this whole time. Furthermore, I extend many thanks to all my family and friends who have cheered me on and supported me throughout this process. You all mean the world to me. Lastly, I would like to acknowledge the Republic of Turkey - Ministry of National Education department for the funding and financial support of me throughout years. vii TABLE OF CONTENTS LIST OF TABLES ....................................................................................................................xii LIST OF FIGURES ................................................................................................................. xiv CHAPTER ONE GENERAL INTRODUCTION ........................................................................1 REFERENCES ...........................................................................................................................7 CHAPTER TWO LITERATURE REVIEW .............................................................................. 11 2.1. INTRODUCTION .............................................................................................................. 12 2.2. BIOLOGICAL AND PHYSICAL CHARACTERIZATION OF CONIFERS ..................... 13 2.2.1. Taxonomical Background of Abies and Pinus and Their Distribution............................... 13 2.2.2. Morphology and Physiology of Fir and Pine Species ....................................................... 17 2.2.3. Morphology and Physiology of Model Species for this Study .......................................... 18 2.2.4. Ecological and Economic Importance of Conifer Species ................................................ 21 2.3. TREE RESPONSES TO ABIOTIC STRESSES ................................................................. 23 2.3.1. Water or drought stress .................................................................................................... 23 2.3.2. Heat stress ....................................................................................................................... 25 2.3.3. Cold Stress (Chilling and Freezing) ................................................................................. 27 2.3.4. Salinity stress .................................................................................................................. 28 2.4. PLANT WATER STRESS DRIVING FORCES ................................................................ 29 2.5. PLANT WATER STRESS RESPONSE MECHANISMS .................................................. 31 2.6. PLANT RESPONSES TO WATER STRESS ..................................................................... 33 2.6.1. Morphological responses ................................................................................................. 33 2.6.1.1. Growth and development ...................................................................................... 33 2.6.2. Physiological responses ................................................................................................... 34 2.6.2.1 Root signaling ........................................................................................................ 35 2.6.2.2. Photosynthesis ...................................................................................................... 36 2.6.2.3. Stomatal activities ................................................................................................. 36 2.6.2.4. Chlorophyll content .............................................................................................. 37 2.6.2.5. Tissue water relations............................................................................................ 37 2.6.2.6. Osmolyte accumulation and adjustment ................................................................ 38 2.6.2.7. Abscisic acid (ABA) ............................................................................................. 39 2.7. ASSESMENT METHODS FOR WATER STRESS IN TREES ......................................... 39 2.7.1. Plant-based Assessments
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