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Effects of Ozone and Water Stress on Plant Growth and Physiology

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Effects of Ozone and Water Stress on Plant Growth and Physiology Effects of Ozone and Water Stress on Plant Growth and Physiology by Sarah Rosemary Davidson This thesis is submitted for the degree of Doctor of Philosophy and for the Diploma of Imperial College Imperial College of Science, Technology and Medicine University of London July 1990 1 Abstract The aim of this project was to investigate the combined effects of ozone and water stress on the growth and physiology of V id a fa b and a Fagus sylvatica. In a series of experiments V i d a fa b was a exposed to ozone for one week, and to water stress for two weeks. Exposure to ozone either preceded, or coincided with, the first day of exposure to water stress. Exposure to ozone resulted in increased leaf conductance, and in some experiments, stimulated shoot growth, although there was no effect of ozone on rootrshoot partitioning. Water stress either had no effect on plant responses to ozone, or reduced the positive impact of ozone. The timing of exposure to ozone and water stress influenced the degree of visible ozone injury and the occurrence of ozone/water stress interactions on gas exchange, but not growth. From June to September 1988, Fagus sylvatica saplings were exposed to episodes of ozone and simultaneous water stress followed by ’recovery’ periods. This study was designed to determine dose-response relationships over a range of ozone concentrations typical of different British summers. Ozone-induced increases in leaf conductance and photosynthesis occurred only in water stressed plants. Root weight measured in the subsequent spring was reduced by ozone in well watered plants, but increased by ozone in water stressed plants. Therefore ozone reduced the negative impact of water stress on root weight. Ozone at concentrations typical of British summers can affect the growth and gas exchange of V i d a fa b aand Fagus sylvatica, and can affect plant sensitivity to water stress. In both species ozone consistently had a delayed impact on gas exchange. Plant responses to ozone and water stress appear to be dependant on the concentration of ozone, the severity of water stress, and the duration of the two stresses. 2 Acknowledgements Firstly, I would like to thank my supervisors Dr Mike Ashmore and Professor Nigel Bell for their advice and guidance at all stages of this research and their encouragement especially over the last few months. I am indebted to three other people; Cathy Garretty, for her unfailing support and encouragement in all things (beans, beech and banana sandwiches!); Mike Holland, for being a patient and generous teacher of many essential thesis-writing skills; and Nigel Ainsworth (A.H.) for sound advice and general aggro. My special thanks go to everyone in the Air Pollution Group at Silwood for, amongst many other things, being such good company. I could not have wished for better work mates. I am most grateful to my family for their advice, constant support and encouragement and for Elenor, who has proved invaluable. Talking of invaluable, Liam deserves a special mention for his enthusiasm, help and encouragement in the preparation of this thesis. Finally, I must thank all my dear friends at Silwood, especially my flat mates past and present (Carol, Guy and Simon, Katy, Paul and Mike). This study was carried out under the tenure of a Natural Environment Research Council Studentship. 3 Table of Contents Pape Abstract 2 Acknowledgements 3 Table of Contents 4 List of Figures 10 List of Tables 15 List of Plates 19 Chapter 1 Introduction 20 1 Ozone Pollution 21 1.1. The formation of ozone 21 1.2. Ozone concentrations in the United Kingdom 22 2 Ozone and plant water stress 22 3 Plant responses to ozone 23 3.1. Ozone and plant gas exchange 23 3.1.1. Stomatal conductance 24 3.1.2. Photosynthesis 25 3.2. The effect of ozone on plant cuticles 26 3.3. Ozone and plant growth 28 3.4. The effect of ozone on plant growth regulators 29 4 4 Combined effects of ozone and water stress 30 4.1. Ozone/water stress studies on crops: the NCLAN program 31 4.2. Ozone/water stress studies on trees 35 5 The aims of this project 36 Chapter 2 Effects of ozone and water stress on Vicia faba 38 1 Introduction 39 2 Materials and methods 41 2.1. Description of experiments 41 2.1.1 Ozone and control treatments 41 2.2. Greenhouse fumigation system 41 2.2.1. Ozone generation 41 2.2.2. Ozone monitoring 44 2.3. Greenhouse conditions 44 2.4. Plants 44 2.5. Soil water stress 45 2.5.1 Control and measurement of soil water deficit 45 2.5.2. Measurement of soil water potential (experiment 4) 47 2.6. Plant water stress (experiment 4) 47 2.6.1. Measurment of leaf water potential 47 2.6.2. Measurement of relative leaf water content 48 2.7. Measurment of leaf conductance: porometry 48 2.8. Gas exchange measurements 49 2.9. Destructive harvests 53 2.10. Non destructive growth measurements and visible injury assessments 53 3 Data analysis 55 3.1. Ozone concentration: variation between chambers 55 3.2. Soil water stress 56 3.2.1. Soil water deficit 56 5 3.2.2. Soil water potential 56 3.3. Plant water stress 56 3.4. Growth data 56 3.5. Gas exchange 57 3.5.1. Leaf conductance 57 3.5.2. Photosynthesis 57 3.6. Leaf injury 57 4 Results 58 4.1. Greenhouse conditions 58 4.2. Ozone exposures 58 4.3. Soil water stress 63 4.3.1. Soil water deficit 63 4.3.2. Soil water potential 63 4.4. Plant water stress 63 4.4.1. The effect of water stress 68 4.4.2. The effect of ozone on leaf water status 68 4.5. Visible ozone injury 68 4.5.1 The effect of water stress 70 4.5.2. Leaf injury in experiment 4 70 4.6. Plant growth 70 4.6.1. Intermediate harvests 70 4.6.2. Final harvests 72 4.6.2.1. The effect of water stress 72 4.6.2.2. The effect of ozone and ozone/water stress interactions 81 4.7. Gas exchange 88 4.7.1. Leaf conductance 88 4.7.1.1. The effect of water stress 88 4.7.1.2. The effect of ozone and ozone/water stress interactions 88 4.7.2. Photosynthesis 102 4.7.2.1. The effect of water stress 102 4.7.2.2. The effect of ozone and ozone/water stress interactions 110 6 5 Discussion 112 5.1. Ozone fumigation 112 5.2. Water stress 112 5.3. Growth 112 5.4. Gas exchange 117 5.5. Visible ozone injury 123 Chapter 3 Effects of ozone and water stress on Faeus svlvatica 125 1 Introduction 126 2 Materials and methods 128 2.1. Experimental design 128 2.2. Ozone exposures 128 2.3. Fumigation system 133 2.3.1. Ozone generation 133 2.3.2. Ozone monitoring 135 2.4. Plants 135 2.5. Soil w ater stress 136 2.6. Plant w ater stress 136 2.6.1. Measurement of leaf water potential 136 2.6.2. Measurement of leaf water content 137 2.7. Gas exchange measurements 137 2.8. Growth measurments 138 2.8.1. Non destructive 138 2.8.2. Destructive harvests 139 2.9. Visual assessments 139 2.10. Aphid control 139 2.11. Autum n leaf fall 140 2.12. Spring bud burst 140 3 Data analysis 141 3.1. Soil water stress 141 3.2. Plant water stress 141 3.3. Gas exchange measurements 144 7 3.4. Growth measurements 144 3.5. Visual assessments 145 3.6. Autumn leaf fall 145 3.7. Spring bud burst 145 4 Results 146 4.1. Ozone exposures 146 4.2. Soil water stress 146 4.3. Plant water relations 146 4.3.1. Relative leaf water content 146 4.3.1.1. The effect of water stress 157 4.3.1.2. The effect of ozone and ozone/water stress interactions 157 4.3.2. Water stressed plants: the effect of ozone 157 4.4. Plant gas exchange 163 4.4.1. Effects of water stress 163 4.4.2. Effects of ozone 163 4.4.3. Interactions between ozone and water stress 176 4.4.4. Gas exchange measurements in 1989 182 4.5. Plant growth 182 4.5.1. Non destructive growth measurements in 1988 and 1989 182 4.5.1.1. The effect of water stress 182 4.5.1.2. The effect of ozone and ozone/water stress interactions 189 4.5.2. Specific leaf area ratio 196 4.5.3. Destructive harvests 196 4.5.3.1. The effect of water stress 196 4.5.3.2. Ozone/water stress interactions 200 4.6. Visual assessments 205 4.6.1. Characterisation of damage symptoms 205 4.6.2. Assessment of tree damage 205 4.6.2.1. The effect of water stress 207 4.6.2.2. The effect of ozone and ozone/water stress interactions 207 4.7. Autumn leaf loss 214 4.7.1. The effect of water stress 214 8 4.7.2. The effect of ozone 214 4.8. Spring bud burst 218 4.8.1. The effect of water stress 218 4.8.2. The effect of ozone 218 5 Discussion 221 5.1. Ozone fumigation 221 5.2. W ater stress 221 5.3. Gas exchange 225 5.4. Tree health 232 5.5. Growth 233 Chapter 4 Discussion 240 1 The effect of ozone on gas exchange and growth 241 2 The effect of water stress in modifying plant response to ozone 243 3 Ozone and crop yield 248 4 Forest decline 249 References 253 9 List of Figures Page Figure 2.1.
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