Associative and Endophytic Nitrogen-fixing Bacteria and Cyanobacterial Associations

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1 Associative and Endophytic Nitrogen-fixing Bacteria and Cyanobacterial Associations Edited by Claudine Elmerich Institut Pasteur, Paris, France and William E. Newton Department of Biochemistry Virginia Polytechnic Institute and State University Blacksburg, Virginia, U.S.A. 4y Springer

2 TABLE OF CONTENTS Preface to the Series Preface List of Contributors Dedication - Johanna Dobereiner ix xiii xvii xxi Chapter 1. Historical Perspective: From Bacterization to Endophytes C. Elmerich 1 1. The Nitrogen Cycle: Heritage from the 19 th Century 1 2. Nutritional Interactions between Bacteria and Plants 3 3. Associative Nitrogen-fixing Bacteria 8 4. Discovery of Nitrogen-fixing Endophytes Cyanobacterial Associations Concluding Remarks 15 Acknowledgement 16 References 16 Chapter 2. Molecular Phylogeny and Ecology of Root Associated Diazotrophic a- and (3-Proteobacteria M. Schmid and A. Hartmann Introduction Tools for Molecular Phylogeny and in situ Localization of Bacterial Isolates and Communities Molecular Phylogeny and Ecology of Azospirillum and Other Nitrogen-fixing a-subclass Proteobacteria Molecular Phylogeny and Ecology of Herbaspirillum, Diazotrophic Burkholderia spp., and Other N 2 -fixing P-Proteobacteria Conclusions and Prospects for Future Studies 35 Acknowledgements 36 References 36 Chapter 3. Regulation of Nitrogen Fixation and Ammonium Assimilation in Associative and Endophytic Nitrogen fixing Bacteria F. O. Pedrosa and C. Elmerich Introduction Rhizospheric and Endophytic Bacteria: General Features Structural Organization of m/genes Identification of RpoN and Its Involvement in Nitrogen Fixation The Ntr System and the Control of Nitrogen Metabolism and Nitrogen Fixation Regulation of Nitrogen Fixation Conclusions 64

3 vi TABLE OF CONTENTS Acknowledgements 64 References 65 Chapter 4. Chemotaxis in Soil Diazotrophs: Survival and Adaptative Response G. Alexandre and 1. B. Zhulin Introduction Gene-Expression Regulation and Chemotaxis as Adaptive Responses to Environmental Changes Molecular Mechanism of the Chemotactic Response: Learning from Escherichia coli Directed Motility in Soil Diazotrophs Future Studies 82 References 82 Chapter 5. Molecular Genetics of Rhizosphere and Plant-Root Colonization E. Vanbleu andj. Vanderleyden Introduction Motility of Associative Diazotrophs Attachment to Plant Roots Rhizosphere Competence Conclusions 103 Acknowledgements 105 References 105 Chapter 6. Microbial Production of Plant Hormones B. E. Baca and C. Elmerich Discovery of Phytohormones Production and Role of Phytohormones Pathways for Plant Hormone Biosynthesis: Common Routes in Plants, Bacteria and Fungi Major Routes for IAA synthesis in Pathogenic and Beneficial Nitrogen-fixing Bacteria Associated with Plants Multiple Routes for IAA Synthesis in Azospirillum Other Phytohormones Produced by Plant Pathogenic and Nitrogen-fixing Associated and Endophytic Bacteria Plant Growth Promotion (PGP): Role of Bacterial Phytohormone Production, ACC-Deaminase, and Use of Synthetic Auxins Concluding Remarks 136 Acknowledgements 137 References 137 Chapter 7. The Plant Growth-Promoting Effect and Plant Responses S. Dobbelaere and Y. Okon N 2 Fixation vs. "Hormonal" Effects: Historical Perspectives Effects of Azospirillum and Other Diazotrophs on Root Morphology 146

4 TABLE OF CONTENTS vii 3. Effects on Root Function Effects on Plant Growth Future Studies 163 References 164 Chapter 8. Biocontrol of Plant Diseases by Associative and Endophytic Nitrogen-fixing Bacteria R. Bally and C. Elmerich Beneficial Plant-Associated Nitrogen-fixing Bacteria and Biocontrol of Plant Disease Interactions within Microbial Communities: Competition Mechanisms Biological Control against Soil-Borne Diseases Regulation of Biocontrol Properties and Cell-Cell Communication Plant Response to Pathogens and Biological Control in the Rhizosphere Concluding Remarks 183 Acknowledgements 184 References 184 Chapter 9. Endophytic Associations of Azoarcus spp. B. Reinhold-Hurek and T. Hurek Introduction The Rise of Interest in Diazotrophic Endophytes Azoarcus spp. and related Genera: Strictly Plant-Associated versus Soil Bacteria Habitats and Ecophysiology Interactions with Fungi Infection of Roots by Endophytic Diazotrophs: An Active, Specific Process? Concluding Remarks 207 References 207 Chapter 10. Biological Nitrogen Fixation in Sugarcane V. Reis, S. Lee and C. Kennedy Short Story of the Sugarcane-Cropping System Nitrogen-fixing Bacteria Colonising Sugarcane: New Phylogenetic Data, Properties, and Endophytic Status Contribution of BNF to the Sugarcane Crop Effect of N Fertilization on BNF Genes for Nitrogen Fixation and Their Regulation in G. diazotrophicus and H. seropedicae Is Indole Acetic Acid Production an Important Factor in the Ability of G. diazotrophicus to Enhance Growth of Sugarcane? Concluding Remarks 226

5 viii TABLE OF CONTENTS Acknowledgement 227 References 228 Chapter 11. Heterocyst Differentiation and Nitrogen Fixation in Cyanobacteria R. Haselkorn Early History of the Association of Nitrogen Fixation with Heterocysts Cyanobacterial Nitrogenase and nif-gex\ss Organization Pathway of N Assimilation Carbon Metabolism in Heterocysts Genetic Tools for Studying Cyanobacterial Nitrogen Fixation Regulatory Genes Required for Heterocyst Differentiation Prospects 251 Acknowledgement 252 References 252 Chapter 12. Cyanobacterial Associations B. Bergman, A. N. Rai, and U. Rasmussen Introduction Historical Aspects and Landmarks Symbioses with Diatoms (Algae) Symbioses with Fungi Symbiosis with Bryophytes Symbiosis with Pteridophytes Symbioses with Cycads Symbiosis with Gunnera Creation of New Symbioses and Prospects 290 Acknowledgements 292 References 292 Chapter 13. Prospects for Significant Nitrogen Fixation in Grasses from Bacterial Endophytes E. W. Triplett Ultimate Objective of Nitrogen-fixation Research - Nitrogen Fixation in Maize, Wheat, and Rice Understanding the Basic Biology of Endophytic Colonization: Using Klebsiella pneumoniae 342 (Kp342) as the Model Diazotrophic Endophyte Attributes Needed for a Model Diazotrophic Endophyte Future Work Needed to Replace Nitrogen Fertilizer with Diazotrophic Endophytes 310 References 312 Subject Index 315

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