Inorganic Plant Nutrition

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1 Q Inorganic Plant Nutrition Edited by A. Lauchli and R.L. Bieleski Contributors C.J.Asher L.Beevers R.T.Besford R.L. Bieleski P.Boger E.G.Bollard H.Bothe D.Bouma G.D.Bowen EC. Cannon C.C. Delwiche J. Dobereiner W.M. Dugger D.G.Edwards I.B.Ferguson T.J.Flowers R.C.Foster W.H. Gabelman G.C. Gerloff R.H. Hageman A. Lauchli U.Liittge D.Marme H.Marschner J.Moorby M.G.Pitman A. Pollard A.Quispel A.D.Robson R.Roth A.D.Rovira G.Sandmann J.A.Schiff W.R. Ullrich D. Werner R.G.Wyn Jones M.G.Yates With a Foreword by E. Epstein With 131 Figures Springer-Verlag Berlin Heidelberg NewYork Tokyo 1983

2 Contents Part A Introduction A. LAUCHLI and R.L. BIELESKI I. General Chapters of Inorganic Plant Nutrition I.I General Introduction to the Mineral Nutrition of Plants H. MARSCHNER (With 11 Figures) 1 Introduction and Historical Resume Essential Mineral Elements - Plant Nutrients Function of Essential Mineral Elements Beneficial Mineral Elements Recent Developments Calcium Potassium Phosphorus Nitrogen Copper Chlorine 15 2 Uptake and Long-Distance Transport of Mineral Elements Ion Concentration at the Root Surface, Role of the "Rhizosphere" Long-Distance Transport in the Xylem From the Roots to the Shoot Into Fruits, Seeds and Storage Organs Retranslocation of Mineral Elements from Leaves 20 3 Calcium Nutrition of Higher Plants Introduction Calcium Demand of Higher Plants Calcium Uptake by the Roots Long-Distance Transport of Calcium Xylem Transport Phloem Transport Xylem Versus Phloem Transport Role of Phytohormones and Growth Regulators Conclusion and Outlook 29 4 Mineral Nutrition and Physiology of Yield Formation - Sink-Source Relationship Introduction Effect of Mineral Nutrition on Phytohormone Level and Sink Formation Effect of Mineral Nutrients on Fertilization Source-Sink Interactions in Relation to Mineral Nutrition 34 5 Environmental Aspects of Mineral Nutrition Introduction Nitrogen Heavy Metals 39

3 XII Contents Part A 5.2 Heavy Metal Toxicity Heavy Metals in the Food Chain Heavy Metals in the Soil/Plant System Content of Soils Soil Factors Affecting Heavy Metal Accumulation in Plants Genotypic Differences in Heavy Metal Uptake Distribution Within the Plants and Their Organs Heavy Metal Tolerance Concluding Remarks 49 References The Significance of Rhizosphere Micro flora and Mycorrhizas in Plant Nutrition A.D. ROVIRA, G.D. BOWEN, and R.C. FOSTER (With 7 Figures) 1 Introduction : 61 2 Energy Supplies in the Rhizosphere Exudates Secretions Plant Mucilages Mucigel Lysates 63 3 Microbiology of the Rhizosphere Populations of Micro-Organisms Colonization of Roots by Micro-Organisms 64 4 Mathematical Modelling of the Rhizosphere 65 5 Microscopy of the Rhizosphere Light Microscopy Scanning Electron Microscopy (S.E.M.) Transmission Electron Microscopy (T.E.M.) General Description : Origin and Fine Structure of Root Mucilage Microbial Invasion of the Mucilage and the Formation of Mucigel Functions of Root Mucilage and Mucigel The Outer Rhizosphere Invasion of the Root by Microorganisms 73 6 The Role of Rhizosphere Microorganisms in Plant Nutrition Availability of Nutrients ; Nutrient Release and Immobilization Nitrification and Denitrification Nitrogen Fixation Phosphate Availability Minor Nutrients Growth and Morphology of Roots Root Length and Root Hairs Proteoid Roots Nutrient Uptake Processes Physiology and Development 76 7 Mycorrhizas Plant Responses to Infection Mechanisms of the Response Nutrient Availability Absorption Characteristics of the Root Absorption by the Fungus Component Energy Requirements of Mycorrhizas Overview of Mycorrhizas 83

4 Contents Part A XIII 8 General Conclusions 84 References Modern Solution Culture Techniques C.J. ASHER and D.G. EDWARDS (With 3 Figures) 1 Major Differences Between Solution Culture and Soil Culture Mechanical Support Spatial Variation in Root Environment Parameters Temporal Variation in Root Environment Parameters Nutrient Depletion ph Shifts Root-Microorganism Interactions 98 2 Uses and Limitations of Existing Solution Culture Methods Non-Renewed or Intermittently Renewed Water Cultures and Sand Cultures Use in Teaching, Demonstration, and Diagnosis Production of Roots for Ion Transport Studies Nutrient Essentiality Effects of Root Environment Parameters Establishment of Critical Tissue Concentrations Control of Plant Nutrient Status Study of Symbiotic Associations with Microorganisms Commercial Crop Production Mist Culture Flowing Solution Culture The Flow Rate Problem Composition of Flowing Culture Solutions Research Applications Ill Likely Future Developments Commercial Crop Production Summary and Conclusions 115 References Diagnosis of Mineral Deficiencies Using Plant Tests D. BOUMA (With 5 Figures) 1 Introduction Plant Analysis : Physiological Basis Choice of Tissue Factors Affecting the Relationship Between Nutrient Concentration and Yield ' Plant Development Effects of Changes in Age of Tissue Plant Age and Critical Levels Interactions Between Nutrient Elements Environmental Factors Other Factors Affecting Nutrient Composition Physiological and Biochemical Approaches to Diagnosis Introductory Remarks Physiological Approaches Physiological Assessment Nutrient Stress Approaches Based on Photosynthesis Other Approaches 133

5 XIV Contents Part A 3.3 Biochemical Approaches Nitrogen and Molybdenum Phosphorus Potassium and Magnesium Iron and Manganese Copper Zinc Prospects for the Future 140 References Interactions Between Nutrients in Higher Plants A.D. ROBSON and M.G. PITMAN (With 9 Figures) 1 Introduction Interactions Between Nutrients in Monoculture Interactions Between Nutrients Affecting the Absorption of Nutrients Interactions Occurring in the Soil Absorption from Solution at the Root Surface Interactions Between Nutrients Affecting the Utilization of Nutrients Within the Plant Distribution Function Complex Interactions Between Nutrients Involving Several Processes Calcium/Aluminium/Phosphate Zinc/Phosphate Interactions Between Nutrients in Mixed Communities Conclusion 173 References 173 I;6 Import and Export of Mineral Nutrients in Plant Roots U. LUTTGE (With 10 Figures) 1 Introduction: The Dual Role of Roots in the Evolution of Higher Land Plants Relations Between Structure and Transport Functions Along the Length of Roots The Phenomenon of Variations in Transport Functions Along the Length of Roots Structure-Function Relations in Various Root Zones The Root Surface The Cortex } The Endodermis The Stele Variations of Physiological Activities Along the Length of Roots Growth, Differentiation and Hormonal Gradients Bioelectrical Fields Along Roots Differences in Ion Transport Mechanisms Along Roots Root-Shoot Interactions and Circulation in the Whole Plant Some Examples Illustrating General Aspects of Circulation Nitrogen, Sulphur and Phosphorus Conclusion 204 References Cycling of Elements in the Biosphere C.C. DELWICHE (With 5 Figures) 1 The Sources of Plant Constituents Soil and Atmospheric Sources The Weathering Process 212

6 Contents Part A XV 2 The Nature of Cycles The Hydrologic Cycle The Sedimentary Cycle The Magmatic Cycle ' The Geobiological Cycles The Nitrogen Cycle Overall Cycle Features Nitrification Denitrification Nitrogen Fixation Human Influences The Sulfur Cycle Comparison with the Nitrogen Cycle Microbial Oxidation Sulfate Reduction Patterns of Sulfur Movement Human Influences The Phosphorus Cycle Oxidation and Reduction Movement and Transport in the Biosphere Human Influences Other Elements Biological Cycling The Special Significance of Iron and Aluminum Hydrogen Ion Characteristics of Sediments Passive Cycling Possibilities of Deficiency "Open" Versus "Closed" Agricultural Systems 236 References 237 II. Inorganic Nitrogen Nutrition II. 1 Physiology, Biochemistry and Genetics of Dinitrogen Fixation H. BOTHE, M.G. YATES, and F.C. CANNON (With 3 Figures) 1 The Nitrogen-Fixing Organisms and the Nitrogenase Reactions Introduction Nitrogen Fixation by Free-Living Organisms Symbiotic Nitrogen Fixation ' Substrates of Nitrogenase Biochemistry of Nitrogen Fixation Introduction Nomenclature of Nitrogenase Proteins Physicochemical Properties of Nitrogenase Proteins Metal Clusters in Nitrogenase Proteins EPR and Mossbauer Spectroscopy on the MoFe Protein The FeMo Cofactor and the Fe Protein Nitrogenase Proteins in Photosynthetic Organisms The Mechanism of Nitrogenase Activity The Roles of the Two Proteins Evidence for Interaction of MgATP and MgADP with the MoFe Protein The Nature of the Active Site(s) Pathways of Nj-Reduction Electron Transport to Nitrogenase Introduction Ferredoxins 260

7 XVI Contents Part A 3.3 Flayodoxins Electron Donors Mechanisms to Protect Nitrogenase Against Damage by Oxygen In Free-Living Organisms The Heterocysts of Blue-Green Algae The Role of Leghaemoglobin in Legume Nodules Regulation of Nitrogenase Activity and Biosynthesis : Regulation of Nitrogenase Biosynthesis Regulation of Nitrogenase Activity The Hydrogenase-Nitrogenase Relationship The Molecular and Genetic Characterization of Nitrogen Fixation Genes Introduction Them/Genes nif Gene Products Cloning of K. pneumoniae nif Genes A Physical Map of nif Genes Interspecies Homology of Nitrogenase Genes 276 References '. 276 II.2 Dinitrogen-Fixing Symbioses with Legumes, Non-Legume Angiosperms and Associative Symbioses A. QUISPEL (With 7 Figures) 1 Introduction Description of the Main Symbiotic Dinitrogen-Fixing Systems Associative Symbioses Symbioses with Cyanobacteria Distribution Description and Development N 2 Fixation (C 2 H? Reduction) Root Nodules with Actinomycetes: Actinorhizas Distribution Description Infection and Development N 2 Fixation (C 2 H 2 Reduction) ' Leguminous Root Nodules with Rhizobium Distribution Description Infection and Nodule Development ', N 2 Fixation (C 2 H 2 Reduction) Non-Leguminous Root Nodules with Rhizobium The Dinitrogen-Fixing Micro-Symbionts: Isolates and Cultures Introduction Cyanobacteria Frankia, the Endophyte from the Actinorhizas Isolation and Cultivation Specificity Nutrient Requirements Metabolic Activities Rhizobium Isolation and Description Taxonomy Metabolism N 2 Fixation (C 2 H 2 Reduction) Genetics Symbiotic Relations Chemotaxis and Rhizosphere Accumulation Binding of Rhizobium to Root Hairs 305

8 Contents Part A XVII 4.3 Root Hair Deformation and Infection-Thread Formation Cell Wall Degrading Enzymes The Role of Plant Hormones in Nodule.Formation Miscellaneous Problems The N 2 -Fixing System Introduction Bacteroids The Bacteroid-Containing Plant Cells Nitrogenase NH 3 Assimilation Oxygen Regulation and Leghaemoglobin Hydrogen Production and Hydrogen Uptake Root Nodules as Part of the Whole Plant Concluding Remarks 323 References Dinitrogen Fixation in Rhizosphere and Phyllosphere Associations J. DOBEREINER (With 2 Figures) 1 Introduction Characterization of Rhizocoenoses Sugar Cane - Beijerinckia ' Paspalum notatum - Azotobacter paspali Azospirillum Rhizocoenoses Taxonomy of Azospirillum spp Root Infection Host Plant Specificity Physiology of Azospirillum Associations with Other N 2 -Fixing Bacteria Agronomic Aspects Plant Genotype Effects Environmental Effects Inoculation Phyllosphere Associations Microorganisms in the Phyllosphere Nitrogen Fixation in the Phyllosphere General Conclusion 344 References. : Uptake and Reduction of Nitrate: Bacteria and Higher Plants L. BEEVERS and R.H. HAGEMAN 1 Introduction Available Nitrogen Sources Species Differences in Ammonium and Nitrate Utilization Influence of Ammonium or Nitrate on Cation Uptake Nitrate Uptake 354, 2.4 Influence of Ammonium on Nitrate Uptake and Utilization Nitrate Reduction Bacteria Dissimilatory Nitrate Reductase Assimilatory Nitrate Reduction in Bacteria Characterization of Nitrate Reductase from Higher Plants Molybdenum in Nitrate Reduction Nitrite Reduction Assimilatory Bacteria Dissimilatory Bacteria Nitrite Reductase in Plants 361

9 XVIII Contents Part A 6 Location of Enzymes of Nitrate Assimilation in Higher Plants Provision of Reductant for Nitrate Assimilation in Higher Plants Regulation of Nitrate Reductase in Higher Plants Substrate Hormonal Molybdenum Ammonium Light Genetic In Vivo Controls Concluding Thoughts 368 References 369 II.5 Uptake and Reduction of Nitrate: Algae and Fungi W.R. ULLRICH (With 4 Figures) 1 Introduction Nitrate and Nitrite Reduction in Algae Nitrate Reductase of Eucaryotic Algae Nitrate Reductase in Blue-Green Algae Nitrite Reductase in Algae Location of Nitrate and Nitrite Reduction in Algal Cells Stoichiometry Between Nitrate Reduction and O 2 Exchange Nitrate Uptake in Algae General Remarks Substrate Affinity Light Dependence ph-dependence Dependence on Carbon Sources Inhibition by Anions Inhibition by Ammonia and Amino Compounds Effect of Metabolic Inhibitors and Uncouplers Stoichiometry Between the Uptake of Nitrate and that of Other Ions Transport Mechanism Nitrite Uptake in Algae General Remarks on Regulation of Nitrate and Nitrite Uptake Uptake and Reduction of Nitrate and Nitrite in Fungi 391 References 393 III. Metabolism of Sulfur and Phosphorus III.l Reduction and Other Metabolic Reactions of Sulfate J.A. SCHIFF (With 6 Figures) 1 Introduction The Place of Sulfate Reduction in the Sulfur Cycle Phylogenetic Distribution of Reactions Involving Sulfate Transfer and Reduction Sulfate Uptake, Activation and Transfer Sulfate Reduction Detailed Reactions of the Two Assimilatory Pathways The APS Pathway The PAPS Pathway Location of Sulfate Reduction in Tissues and Organs of Multicellular Plants 413

10 Contents Part A XIX 6 Speculations on the Origin and Evolution of Pathways of Sulfate Reduction 413 References, 416 III.2 Physiology and Metabolism of Phosphate and Its Compounds R.L. BIELESKI and I.B. FERGUSON (With 4 Figures) 1 Introduction Uptake and Transport of Phosphate Efflux of Phosphate, and Aspects of Phosphate Deficiency Phosphorus Compartments and Pools The Form of Phosphorus in the Cell Synthesis and Turnover of Phosphorus Compounds Dynamics of Phosphate Use in the Plant Conclusions 445 References 445 Author- and Subject Index (see Part B)

Inorganic Plant Nutrition

Inorganic Plant Nutrition Edited by A. Lauchli and R.L. Bieleski Contributors C.J.Asher L.Beevers R.T.Besford R.L. Bieleski P.Boger E.G. Bollard H.Bothe D.Bouma G.D.Bowen F.C. Cannon C.C.Delwiche J. Dobereiner