Manganese in Soils and Plants

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Manganese in Soils and Plants Proceedings of the International Symposium on 'Manganese in Soils and Plants' held at the Waite Agricultural Research Institute, The University of Adelaide, Glen Osmond, South Australia, August 22-26, 1988 as an Australian Bicentennial Event Edited by Robin D. Graham Department of Agronomy, Waite Agricultural Research Institute, University of Adelaide, Glen Osmond, Australia Robert J. Hannam Northfield Laboratories, South Australian Department of Agriculture, Adelaide, Australia and Nicholas C. Uren School of Agriculture, La Trobe University, Bundoora, Australia Publication sponsored by BHP-UTAH Minerals International { universitatsbibli^t :. HANNOVER TECHNIGCHE fjnfqrmatiqnsriibliqthek KLUWER ACADEMIC PUBLISHERS DORDRECHT / BOSTON / LONDON

Table of Contents 6 List of Contributors Preface Organising Committee Financial Supporters xi xin xv xvii Historical Preface l R.D. Graham and J.P.Quirk 1. An Introduction to Manganese Biological Chemistry N.P. Hughes and R.J.P. Williams 1. Introduction 7 2. Manganese chemistry 7 3. The biological chemistry of Mn 14 4. Summary 18 Manganese in Soils 2. Geochemistry of Manganese in Soil R.J. Gilkes and R.M. McKenzie 1. Introduction 23 2. The crystal chemistry of Mn 2 3 3. Manganese in rock-forming minerals 2 3 4. Manganese in soils 26 5. Conclusions 34 3. Inorganic Reactions of Manganese in Soils W.A. Norvell 1. Introduction 37 2. Solubility relationships of Mn solid phases 3 7 3. Forms of Mn in soil solution 41 4. Surface reactions 4 5 5. Fertilizer reactions 52

VI 4. Manganese Redox Reactions and Organic Interactions in Soils Richmond J. Bartlett 1. Introduction: Mn, key to life 5 9 2. Soil Mn redox methods 5 9 3. phandpe 61 4. Reverse dismutation of Mn 6 2 5. Electron demand and net oxidation studies 6 3 6. ManganeseCm) catalysis and free radicals 6 6 7. Contributions of components of the living soil 6 8 8. Manganese mediation of soil N transformations 70 9. Role of Mn in formation of humic polymers 71 5. The Biology of Manganese Transforming Microorganisms in Soil W.C. Ghiorse 1. Introduction 7 5 2. Ecology 7 5 3. Taxonomy 7 7 4. Physiology and biochemistry 78 5. Conclusions 82 6. Determination of Plant-available Soil Manganese H.M. Reisenauer 1. 2. 3. 4. 5. 6. Introduction Plant factors in soil Mn availability SoilMn Soil tests for plant available Mn Available soil Mn Conclusions 87 87 88 91 92 95 Manganese in Plants 7. The Uptake and Translocation of Manganese by Plant Roots D.T. Clarkson 1. Introduction 101 2. The nature of the supply of Mn to the root surface 101 3. Electrochemical considerations 103

4. Kinetics of Mn 2+ uptake 104 5. Metabolic dependence of Mn 2+ uptake 105 6. Interactions between the uptake and translocation of Mn 2+ and other ions 106 7. Uptake rates compared with growth requirement for Mn 109 8. Concluding remarks 110 Vll 8. Distribution and Movement of Manganese in Plants Jack F. Loneragan 1. Introduction 113 2. Movement of Mn in plants 113 3. Distribution of Mn in plants 117 4. Conclusions 121 9. The Biochemistry of Manganese in Plants James N. Burnell 1. Introduction 125 2. Manganese in proteins 125 3. Manganese as a cofactor 128 4. C 4 photosynthesis 132 5. Concluding remarks 133 10. Physiological Functions of Manganese in Plants Lindsay C. Campbell and Ross O. Nable 1. Introduction 139 2. Growth physiology of Mn 139 3. Environmental interactions with Mn 144 4. Manganese in metabolism 145 5. Future directions 149 11. The Role of Manganese in Disease Resistance D.M. Huber and N.S. Wilhelm 1. Introduction 155 2. Effect of disease on Mn 155 3. Effect of Mn on disease 156 4. Mechanisms of disease control with Mn 164 5. Conclusions 167

Vlll 12. The Physiology of Manganese Toxicity W.J. Horst 1. Introduction 175 2. Manganese toxicity symptoms 175 3. Factors affecting Mn tolerance levels of plant tissues 176 4. Relationships between toxicity symptoms and physiological effects of high Mn concentrations in plant tissues 178 5. Physiology of differential Mn tolerance 183 6. Summary and conclusions 184 Manganese in Soil-Plant Systems 13. Mechanisms of Manganese Acquisition by Roots from Soils H. Marschner 1. Introduction 191 2. Plant and soil factors in absorption 191 3. Dynamics of Mn at the soil-root interface (rhizosphere) 195 4. Conclusions 201 14. Occurence and Correction of Manganese Deficiency in Plants D.J. Reuter, A.M. Alston and J.D. McFarlane 1. Introduction 205 2. Occurrence of Mn deficiency 205 3. Prevention and correction of Mn deficiency 211 4. Conclusions 218 15. Manganese Fertilizers K.H. Walter 1. Introduction 225 2. The chemistry of Mn sources 225 3. Incorporation of Mn into macro-nutrient fertilizers 232 4. Conclusions 238

16. Detection of Manganese Deficiency and Toxicity in Plants R.J. Hannam and K. Ohki IX 1. Introduction 243 2. Diagnosis of Mn deficiency 243 3. Prognosis of Mn deficiency 251 4. Diagnosis of Mn toxicity 252 5. Conclusions 255 17. Genotypic Differences in Tolerance to Manganese Deficiency R.D. Graham 1. Introduction 261 2. The agronomic case for breeding for Mn efficiency 261 3. Genotypic differences in tolerance to Mn efficiency 263 4. Genetics of Mn efficiency 266 5. Mechanisms of Mn efficiency 268 6. Screening techniques for identifying Mn-efficient genotypes 271 7. Conclusions 273 18. Distribution and Amelioration of Manganese Toxic Soils E. Schlichting and L.A. Sparrow 1. Introduction 277 2. The global distribution of Mn toxic soils 277 3. The development of Mn toxic soils 279 4. The amelioration of Mn toxic soils 284 5. Conclusions 288 19. Genetics and Breeding of Plants Tolerant of Manganese Toxicity CD. Foy, B.J. Scott and J.A. Fisher 1. Introduction 293 2. Examples of differential tolerance 293 3. Breeding for Mn tolerance 294 4. Development and use of Mn tolerant germplasm 296 5. Physiology of differential Mn tolerance 296 7. Conclusions 302

X 20. Techniques for Research on Manganese in Soil-Plant Systems N.C. Uren, C.J. Asher and N.E. Longnecker 1. Introduction 309 2. Solution culture systems 309 3. Seed reserves of Mn 314 4. Manganese oxides as sources of Mn 315 5. Analysis and detection of Mn in soils and plants 317 6. Soil preparation and pot culture 320 7. Fractionation procedures 322 21. Manganese in Soils and Plants An Overview A.D. Robson 1. Introduction 329 2. Reactions of Mn in soils and plants 329 3. Diagnosis and correction of Mn deficiency and toxicity 332 4. Conclusion 333 Glossary of Plant Names 335 Index 337