Emanuel Epstein Arnold J.

Bloom
University of California, Davis University of California, Davis

Sinauer Associates, Inc. Publishers

Sunderland, Massachusetts

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or disseminated in any form without express written permission from the publisher.
Contents

PART I Elements

1 Introduction and History 3

The Scope of Mineral Plant Nutrition 3 Heredity and Environment 5
Mining the Environment 3 The History of Plant Nutrition Research 6
Translocation 4 References 14
Functions of Elements 5

2 The Media of Plant Nutrition 17

The Variety of Nutrient Media 17 The Osmotic Potential of Nutrient
Solutions 33
Soil 18
Culture Solutions Compared with Soil
Roots in Soil 22 Solutions 34
Artificial Media 25 References 36
Nutrient Solutions 30
The pH of Nutrient Solutions 31

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X CONTENTS

3
Water 41
Inorganic Components of Plants 41
Deficiency Symptoms: Individual
Dry Matter 43 Elements 57
B: Boron 57
Mineral Composition 44
Ca: Calcium 57
Essential and Other Mineral Elements 44
Cl: Chlorine 57
Macronutrient Elements 46 Cu: Copper 59
Micronutrient Elements 46 Fe: Iron 59
Comparative Elemental Requirements of K: Potassium 59
Higher Plants 47 Mg: Magnesium 59
Na: Sodium 47 Mn: Manganese 59
Se: Selenium 48 Mo: Molybdenum 61
Co: Cobalt 48 N: Nitrogen 61
Si: Silicon 48 Ni: Nickel 61
Other Elements 49 P: Phosphorus 62
Quantitative Considerations 51 S: Sulfur 62
Deficiencies and Tissue Analysis 51 Zn: Zinc 62
Deficiency Symptoms: General References 63
Discussion 55

PART II Transport

4 Nutrient Absorption by Plants 69

The Need for a Membrane 69 The Structure of Plant Cells 75
Membrane Structure 70 Passive versus Active Transport of
Apoplast and Cell Walls 73 Solutes 75

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 CONTENTS XI

Active Solute Transport into Cells and Energy Coupling 89

Tissue 78 Solute Transporters 92
Algae as Model Systems 79 References 95
Root Solute Transport 82

5 Upward Movement of Water

and Nutrients 99
Cellular versus Long Distance Mechanisms: The CraftsBroyer
Transport 99 Hypothesis and Beyond 118
Mechanisms: The Endodermis, Stele,
Pores, Pipes, and Pathways 100 Stem, and Leaf 120
Tissues and Organs 104 Some Particular Points and Problems
Movement of Water through the 125
Plant 108 Beyond the Leaf 131
Pathway 108 Leaching 131
Mechanisms 110 Guttation 132
Upward Movement of Ions 114 Excretion of Salt 132
Pathway 114 Leaf Fall 132
References 133

6 Downward Movement of Food

and Nutrients 141
Basic Dilemma 141 Phloem Transport of Inorganic Solutes
Again: Pores, Pipes, and Pathways 143 151
Speed of Phloem Transport 144 The Nature of Controversy 156
Direction of Phloem Transport 147 References 157
Pressure Flow Hypothesis 148

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XII CONTENTS

Metabolism
PART III & Growth

7 Nitrogen and Sulfur:

A Tale of Two Nutrients 163
Nitrogen in the Environment 164 Nitrogen Assimilation 180
Little Shop of Horrors 166 Nitrate Assimilation 181
Biological Nitrogen Fixation 167 Ammonium Assimilation 186
Symbiotic Nitrogen Fixation 171 Sulfur in the Environment 189
Courtship between Legumes and Sulfate Assimilation 190
Rhizobia 173 Nitrogen and Sulfur Regulation 192
Mycorrhizal Associations 177 References 193
Ammonium and Nitrate 178

8 Mineral Metabolism 201

Cell Components 201 Manganese 219
Cell wall 201 Molybdenum 220
Vacuole 202 Nickel 220
Cytoplasm 202 Nitrogen 221
Elements 203 Phosphorus 221
Boron 205 Potassium 225
Calcium 207 Silicon 227
Chlorine 210 Sodium 231
Copper 212 Sulfur 233
Iron 213 Zinc 233
Magnesium 217 References 235

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 CONTENTS XIII

9 Nutrition and Growth 243

Nutrition of Seeds 243 Photosynthesis, Primary Productivity, and
Developmental Stages and Root Nutrients 258
Growth 246 Crop Yields and Nutrient Efficiency 260
Root Hairs and Lateral Roots 253 Testing for Nutrients 265
Regulation of Nutrients by Plants 255 References 266
Seasonal Dynamics 257

Heredity &
PART IV Environment

10
Background 277
Physiological Genetics
and Molecular Biology 277
Ammonium Transporters 288
For the Common Good 278 Heavy Metal Transporters 289
Nitrate Transporters 291
Wild versus Domesticated Species 278
Phosphate Transporters 292
Molecular Biology of PlantNutrient
Sulfate Transporters 293
Relations 281
Water Transporters 293
Proton Pumps 281
Other Transporters 294
Potassium Transporters 284
Calcium Channels and Pumps 286 References 295
Sodium Transporters 287

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XIV CONTENTS

11
Concepts 301
Ecology and Environmental Stress 301
Freezing 323
Ecology and Evolution 301 Heat 324
Plant Physiological Ecology 303 Water Stress 325
Phenotypic Plasticity 305 Drought 325
The Concept of Ecotype 308 Flooding 327
Population Biology 309 Mineral Stresses 330
Communities 311 Salinity 332
Mycorrhizal Associations 312 Physiology of Salt Stress 334
Parasitic Plants 316 Aluminum 339
Stress 319 Heavy Metals 340

Temperature Stress 320 References 342

Chilling 320

12 Big Picture: Past, Present, Future 355

A Brief History of the World 355 Predictions 366
The Atmosphere 358 Transporters 366
Global Climate Change 360 Nutrient chemotropism 367
Nutrient assimilation and
Nutrient Cycling 362 sequestration 368
Phosphorus 362 Horticultural and agronomic
Potassium 364 practices 368
Salt 365 Finale 370
References 370

Author Index 375

Subject Index 390

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