A. Sakai W. Larcher - l-o o Frost Survival of Plants Responses and Adaptation to Freezing Stress With 200 Figures Springer-Verlag Berlin Heidelberg New York London Paris Tokyo
1. Low Temperature and Frost as Environmental Factors.... 1 1.1 Low Temperature Hazards 1 1.2 Cold, Frost, and Snow 1 1.3 The Occurrence of Cold, Frost, and Snow 3 1.4 Frost and Snow in the Plant's Environment 9 1.5 Time of Onset, Severity and Duration of Frost: The Freezing Risk 15 1.6 Temperature Fluctuations in Wintering Trees 19 2. The Freezing Process in Plants 21 2.1 Freezing of Water and Aqueous Solutions 21 2.1.1 Ice Nucleation. 21 2.1.2 Freezing of Solutions 23 2.1.3 Sequence of Events in the Freezing of Liquids.... 24. 2.1.4 Vitrification of Water and Solutes 24 2.2 Freezing of Plant Cells 26 2.2.1 Extracellular Freezing 26 2.2.2 Intracellular Freezing 29 2.2.3 Freezing of Intact Tissues 31 2.2.4 Threshold Freezing Temperature and Supercooling of Plant Tissues 33 2.2.5 Initiation and Progress of Freezing in the Whole Plant. 36 3. Freezing Injuries in Plants 39 3.1 Typology of Freezing Mechanisms 39 3.2 Causes of Death by Freezing 41 3.3 Phenomenology of Frost Damage 47 3.3.1 Symptoms of Injury 47 3.3.2 Freezing Damage Peculiar to Trees 48 3.4 Results of Freezing Injury and Chances of Recovery.... 54 4. Mechanisms of Frost Survival 59 4.1 Components of Frost Survival 59 4.1.1 Frost Mitigation 59
VIII 4.1.2 Frost Exclusion 62 4.1.3 Components of Frost Resistance 65 4.2 Supercooling as a Survival Mechanism 70 4.2.1 Persistent Supercooling of Leaves 70 4.2.2 Deep Supercooling of Xylem Ray Parenchyma Cells. 72 4.3 Extraorgan and Extratissue Freezing 76 4.3.1 Deep Supercooling and Extraorgan Freezing of Hydrated Seeds 77 4.3.2 Deep Supercooling and Extraorgan Freezing of Floral Primordia of Woody Angiosperms 81 4.3.3 Extraorgan Freezing of Conifer Buds 85 4.4 Comparison and Classification of Frost Survival Mechanisms 89 4.5 Survival at Ultralow Temperatures 91 4.5.1 Desiccated Plant Tissues 91 4.5.2 Hydrated Plant Tissues 93 5. Cold Acclimation in Plants 97 5.1 The Annual Course of Frost Resistance 97 5.2 The Seasonal Cold Acclimation Process 99 5.2.1 Seasonality Timing 100 5.2.2 Induction of Cold Acclimation and the First Stage of the Hardening Process 102 5.2.3 The Second Stage of Frost Hardening 105 5.3 Environmental Control of the Level of Frost Resistance..107 5.3.1 Fluctuation of Frost Resistance During Winter...108 5.3.2 Water Content of the Plant and Frost Resistance...112 5.4 Biochemical and Structural Changes During Cold Acclimation 113 5.4.1 Metabolie Pathway Shifts from the Growing State to the Dormant State 113 5.4.2 Sugars and Related Compounds 114 5.4.3 Amino Acids 117 5.4.4 Nucleic Acids and Proteins 118 5.4.5 Lipid Changes 124 5.4.6 Abscisic Äcid 132 5.4.7 Cytological Changes During Cold Acclimation....133 6. Frost Resistance in Plants 138 6.1 Genetic Variation in Frost Resistance 138 6.1.1 Differences in Frost Resistance Between Plant Taxa. 138 6.1.2 Differences in Frost Resistance Within Populations.. 142 6.1.3 Inheritance of Frost Resistance 153
IX 6.2 Differences in Frost Resistance of Various Plant Organs and Tissues 157 6.2.1 Vegetative Organs 160 6.2.2 Reproductive Organs 167 6.3 Ontogenetic Variation in Frost Resistance 168 6.3.1 Variation of Frost Resistance During the Cell Cycle. 169 6.3.2 Frost Resistance of Seeds 170 6.3.3 Frost Susceptibility During Germination and at the Early Seedling Stage 171 6.3.4 Frost Resistance During Establishment and Ageing.. 172 7. Regional Distribution of Plants and Their Adaptive Responses to Low Temperatures 174 7.1 Woody Plants Within the Tropics 175 7.1.1 Mangroves 179 7.1.2 Palms 181 7.2 Evergreen Broad-Leaved Trees and Shrubs 185 7.2.1 Temperate Rain Forest Species 186 7.2.2 Mediterranean Sclerophylls 196 7.3 Transition from an Evergreen to a Deciduous Flora and Its Consequences for Frost Survival 200 7.3.1 Woody Plants 202 7.3.2 Ferns 206 7.4 Conifers 210 7.4.1 The Freezing Resistance of Various Conifer Families and Its Connection with Their Distribution Range..211 7.4.2 Characteristics of Freezing Resistance of Conifers..213 7.5 Mountain Plants 218 7.5.1 Dwarf Shrubs and Herbs of Temperate Mountains.. 218 7.5.2 Caulescent Giant Rosette Plants of Tropical High Mountains 224 7.6 Trends of Adaptive Improvement of Low Temperature Resistance in Vascular Plants 229 8. Winter Damage as the Result of a Complexity of Constraints. 235 8.1 Winter Desiccation 238 8.2 Damage Due to Ice Encasement and Compact Snow.... 251 8.3 Harmful Effects of Heavy and Long-Lasting Snow Cover.. 252 8.4 Winter Survival: A Complex Response 259
X Appendix 260 Determination of Frost Resistance for Comparative Studies.. 260 Field Survival 260 Artificial Freezing Tests 261 Analysis of the Freezing Process 262 Frost Treatment of the Samples 263 Identification of Frost Injuries 264 Quantification of Frost Resistance 266 References 267 Terminology and Definitions 305 Taxonomic Index 307 Subject Index 317