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Application of Nitrogen-Fixing Systems in Soil Management

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Application of Nitrogen-Fixing Systems in Soil Management FAO SOILS BULLETIN 49 application of nitrogen-fixing systems in soil management F000 AND AGRICULTURE ORGANIZATION OF THE UWITED NATION'S ROW L PREFACE In view of the continuing increase in cost and scarcity of mineral fertilizers resulting from the use of high-cost fossil energy, there is renewed interest in organic recycling and biological nitrogen-fixation to improve soil fertility and productivity. FAO, in cooperation with the Swedish International Development Authority (SIDA) has assumed a leading role in this respect, including the sponsoring of a series of important international meetings to promote the use of organic materials and biofertilizers. Examples of these meetings include: the Expert Consultation on Organic Materials as Fertilizers held in Rome in 1974; the Workshop on Organic Recycling in Asia held in Bangkok in 1976 and the Workshop on Organic Materials and Soil Productivity in the Near East held in Alexandria in 1978. The workshop in Alexandria recommended the further promotion of research, development, application and dissemination of information available on various aspects of biological nitrogen-fixation, including symbiotic systems of rhizobia/legume and Azolla/blue-green algae, and free-living nitrogen-fixing bacteria and blue-green algae. Dr Y.A. Hamdi, a delegate and invited speaker at the workshop was requested to prepare the present Bulletin, which reviews the literature and provides information on the characteristics, production and use of biofertilizers in agriculture. Although some of the methods described are being continually improved upon, it is hoped that the compilation of various aspects of nitrogen-fixation under one cover in this Bulletin will be of interest and assistance not only to research workers, but also to extension workers and planners concerned with the further development and refine- ment of these natural systems for soil improvement and management throughout the various agro-ecological zones of the world. ACKNOWLEDGEMENTS The Food and Agriculture Organization of the United Nations and the author wish to gratefully acknowledge all organizations and individuals who have assisted directly or indirectly in the preparation and processing of this Bulletin. The author also wishes to express his deep appreciation of assistance from all those scientists and colleagues who made their research papers and photographs available for use as required in the Bulletin. More specific acknowledgements of contributions and sources of information are given in the text, supported by a list of references for further reading as required on these important subjects. TABLE OF CONTENTS Page 1. INTRODUCTION 1 1.1 General 1 1.2 Global Status of Nitrogen 1 1.3 Fixed Nitrogen 2 1.4 Fertilizer Nitrogen 3 2. BIOLOGICAL NITROGEN-FIXATION 5 2.1 Introduction 5 2.2 Free-Living Nitrogen-Fixing Organisms 5 2.2.1 Bacteria 5 2.2.2 Blue-green algae 5 2.3 Associative Symbiosis Between Free-Living Diazotrophs and Other Organisms 6 2.3.1 Loose associations 6 2.3.2 Associations with animals 6 2.3.3 Associations involving morphological modification or accommodation 8 2.4 Biochemistry of Biological Nitrogen Fixation 9 2.4.1 Nitrogenase enzyme system 9 2.4.2 Requirements for nitrogen-fixation 9 3. RHIZOBIA-LEGUME SYMBIOSIS 11 3.1 The Microsymbiont.Rhizohium 11 3.1.1 Taxonomy 11 3.1.2 Morphology and cytology of Rhizobium 11 3.1.3 Cultural characteristics of Rhizobium 11 3.1.4 Physiology of 12 3.1.5 Ecology of rhizobia in soil 14 3.2 The Macrosymbiont. The Legumes 14 3.2.1 Cross-inoculation groups 14 3.2.2 Nodules 16 3.3 Genetic Aspects of Nodulation and Nitrogen-Fixation by Legume-Rhizobia Association 22 3.3.1 The macrosymbiont. Legume host 22 3.3.2 The microsymbiont. Rhizobium 22 3.4 Legume Inoculation 24 3.4.1 Strain selection 24 3.4.2 Collection of strains 24 3.4.3 Evaluation of strains 24 3.4.4 Bacteria propagation 25 3.4.5 Carriers of rhizobia 25 3.4.6 Inoculant preparation 28 Legume Inoculation (Cont.) Page 3.4.7 Standards of inoculants 28 3.4.8 Multistrain inoculants 28 3.4.9 Tests for control of inoculants 28 3.4.10 Inoculant application 29 3.4.11 Response to inoculation 31 3.4.12 Factors affecting the success of inoculation 36 3.4.13 Amount of nitrogen fixed by different legumes 40 4. BLUE-GREEN ALGAE 45 4.1 Introduction 45 4.2 Nitrogen-Fixing Blue-Green Algae 45 4.3 Classification 45 4.4 Genetics of Blue-Green Algae 48 4.5 The Site of the Nitrogenase Enzyme in Blue-Green Algae 48 4.6 Isolation and Purification of Blue-Green Algae 48 4.6.1 Unialgal cultures 48 4.6.2 Bacteria-free cultures of algae 49 4.6.3 Tests for purity of algal cultures 49 4.6.4 Methods for testing the ability of blue-green algae to fix nitrogen 50 4.7 Growth Rate of Blue-Green Algae 51 4.8 Ecology and Factors Affecting Nitrogen-Fixing Blue-Green Algae 51 4.8.1 Distribution 51 4.8.2 Nutrients for blue-green algae 51 4.8.3 Light 53 4.8.4 Temperature 54 4.8.5 pH 54 4.8.6 Desiccation 54 4.8.7 Oxidation reduction potential 54 4.8.8 Pesticides 55 4.8.9 Algal pathogens 56 4.8.10 Salinity 56 4.9 Amounts of Nitrogen Fixed by Blue-Green Algae 56 4.9.1 Under laboratory conditions 56 4.9.2 Under field conditions 56 4.10 Transfer of Fixed Nitrogen 58 4.11 Production of Vitamins and Growth Substances by Blue-Green Algae 59 4.12 Mass Cultivation of Blue-Green Algae 59 4.12.1 Japanese technology 59 4.12.2 Indian technology 60 4.12.3 Chinese technology 62 4.12.4 Other methods 62 4.13 Algalization or the Application of Blue-Green Algae to Soil 63 4.13.1 Methods of inoculating blue-green algae 63 4.13.2 Effect of blue-green algae on soil and rice yield 64 4.13.3 Effect of blue-green algae on crops other than rice 73 Page 5. FREE-LIVING NITROGEN-FIXING BACTERIA 75 5.1 Azotobacteriaceae. Taxonomy 75 5.2 Genus Azotobacter 75 5.3 Azotobacterin 75 5.3.1 Historical background 75 5.3.2 Production of azotobacterin 75 5.3.3 Methods of application of azotobacterin 77 5.3.4 Effect of azotobacterin on plant yield 78 5.4 Azotobacter paspali 80 5.4.1 Potentiality of nitrogen-fixation 81 5.4.2 Location of nitrogenase activity 82 5.4.3 Amounts of nitrogen fixed in the field 82 5.5 Beijerinckia 82 5.5.1 Taxonomy 82 5.5.2 Geographic distribution 82 5.5.3 Soil pH 84 5.5.4 Vegetation 84 5.5.5 Significance in soil improvement 84 5.5.6 Efficiency of nitrogen-fixation 84 5.6 Azospirillum 84 5.6.1 Isolation of Spirillum lipoferum 85 5.6.2 Taxonomy 88 5.6.3 Location of A. lipoferum 88 5.6.4 Nitrogen-fixation potential 88 5.6.5 Nitrogen-fixation rates 88 5.6.6 Temperature 88 5.6.7 pH 90 5.6.8 Oxygen 90 5.6.9 Response to inoculation 90 6. AZOLLA 93 6.1 Introduction 93 6.2 Taxonomy 93 6.3 Azolla-Azolla anabaena association 94 6.4 Reproduction 96 6.5 Nitrogen-Fixation 96 6.5.1 Effect of combined nitrogen 99 6.5.2 Nitrogen-fixation efficiency 99 6.5.3 Nitrogen-fixation and conversion factors 99 6.5.4 Release of fixed nitrogen by the algae 99 6.5.5 Photosynthesis of Azolla 99 6.5.6 Growth of Azolla 99 6.6 Total Nitrogen Input Estimates of Azolla 100 6.7 Environmental Factors Affecting Azolla 101 6.7.1 Water 101 6.7.2 Light 101 6.7.3 Temperature 101 6.7.4 pH 102 6.7.5 Season and soil conditions 102 Environmental Factors Affecting Azolla (Cont.) Page 6.7.6 Inoculum density 103 6.7.7 Nutrition 103 6.7.8 Salinity and salt concentration 105 6.7.9 Pests of Azolla 106 6.7.10 Pesticides 106 6.8 Control of Azolla Pests 106 6.9 Fungal Diseases 107 6.10 Role of Azolla in Agriculture1 107 6.10.1 Azolla as green manure 107 6.10.2 Azolla grown in the field 108 6.10.3 Crop response to Azolla 109 6.10.4 Availability of Azolla nitrogen 110 6.10.5 Azolla cultivated under rice plants 111 6.10.6 Place of Azolla in rice rotation in Vietnam 111 6.10.7 Double narrow row method 112 6.10.8 Recommendations for Azolla cultivation 114 6.10.9 Nutrients for Azolla during summer114 6.10.10 Azolla and rice yields 115 6.10.11 Azolla as feed 116 6.10.12 Azolla as weed 117 6.10.13 Azolla as means of improving water quality 117 6.10.14 Other potential uses of Azolla 117 7. NODULATED NON-LEGUMINOUS PLANTS 119 7.1 The Macrosymbiont. The Plants 119 7.2 The Endophyte. The Microsymbionts 120 7.3 Cross Inoculation Groups 122 7.4 Nodules 122 7.5 Extent of Nitrogen-Fixation in some Non-Legumes 124 7.6 Other Non-Legume Nitrogen-Fixing Plants 126 8. SOME COSTS OF BIOFERTILIZERS 127 8.1 Legume Inoculants 127 8.2 Algal Inoculants 127 APPENDIXES A. METHODS OF ASSESSING BIOLOGICAL NITROGEN-FIXATION 129 A.1 Introduction 129 A.2 Kjeldahl Method 129 A.3 15N Method 129 A.4 Acetylene Reduction 129 B. RHIZOBIA 134 B.1 Recognition and Characterization of Rhizobia 134 B.2 Litmus Milk Reaction 135 B.3 Legume Seed Pelleting 135 B.4 Techniques for Quality Control of Legume Inoculants 137 Page C.
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