PLANT SURFACE MICROBIOLOGY Ajit Varma Lynette Abbott Dietrich Werner Rüdiger Hampp (Eds.) Plant Surface Microbiology With 138 Figures,2 in Color 1 23 Professor Dr.Ajit Varma Professor Dr.Dietrich Werner Director FG Zellbiologie und Angewandte Botanik Amity Institiute of Microbial Sciences Philipps Universität Marburg Amity University 35032 Marburg Noida 201303 Germany UP, India email:[email protected] email:[email protected] Professor Dr.Rüdiger Hampp Professor Dr.Lynette Abbott Physiological Ecology ofPlants School ofEarth and Geographical Sciences University ofTübingen The University ofWestern Australia 72116 Tübingen Nedlands WA 6009 Germany Australia email:[email protected] email:[email protected] ISBN 978-3-540-74050-6 Springer-Verlag Berlin Heidelberg New York Library ofCongress Control Number: 2007934913 This work is subject to copyright.All rights are reserved,whether the whole or part ofthe material is concerned, specifically the rights oftranslation,reprinting,reuse ofillustrations,recitation,broadcasting,reproduction on microfilm or in any other way,and storage in data banks.Duplication ofthis publication or parts thereofis permit- ted only under the provisions ofthe German Copyright Law ofSeptember 9,1965,in its current version,and per- missions for use must always be obtained from Springer-Verlag.Violations are liable for prosecution under the German Copyright Law. Springer-Verlag is a part ofSpringer Science+Business Media springer.com © Springer-Verlag Berlin Heidelberg 2004, 2008 The use ofgeneral descriptive names,registered names,trademarks,etc.in this publication does not imply,even in the absence ofa specific statement,that such names are exempt from the relevant protective laws and regulations and therefore free for general use. 5 4 3 2 1 0 – Printed oanc id free paper Preface The complexity ofplant surface microbiology is based on combinations.A large number of microbial species and genera interact with several hundred thou- sand species ofhigher plants.At the same time,they interact with each other. Therefore,this book describes only some very important model interactions which have been studied intensively over the last years.The methods developed for some important groups ofmicroorganisms can be used for a large number ofother less studied interactions and combinations.The pace ofdiscovery has been particularly fast at two poles ofbiological complexity,the molecular events leading to changes in growth and differentiation,as well as the factors regulat- ing the structure and diversity ofnatural populations and communities. The area of plant surfaces is enormous.A single maize plant has a leaf surface of up to 8000 cm2,a single beech tree has a leaf surface of around 4.5 million cm2.The leaf area index (LAI) varies from 0.45 in tundra areas up to 14 in areas with a dense vegetation. Calculated for all plant surfaces above ground,the surface area is more than 200millionkm2.This area is still surpassed by the below ground surface areas of plants,especially those with an extensive root hair system.For a single rye plant,a root hair surface of around 400m2has been calculated.Even if this is an exceptional case,it can be assumed that in many plants the root and root hair surface is ten times larger than the surfaces of the above ground plant parts. This means that more than 2000millionkm2of plant surface is present underground.Taking both figures together,it exceeds the land surface area of the planet Earth of 149 million km2 by more than a factor of 10. This volume summarizes and updates both the state ofknowledge and the- ories and their possible biotechnological applications.It will thus be ofinter- est to a diverse audience ofresearchers and instructors,especially biologists, biochemists, agronomists, foresters, horticulturists, mycologists, soil scien- tists, ecologists, plant physiologists, plant molecular biologists, geneticists, and microbiologists. In the planning ofthe book,invitations for contributions were extended to leading international scientists working in the field ofplant surface microbi- VI Preface ology. The basic concepts in plant surface microbiology are discussed at length in 30 chapters including a few specialized and innovative methodolo- gies and novel techniques.The editors would like to express deep appreciation to each contributor for his/her work,patience and attention to detail during the entire production process.It is hoped that their reviews,interpretations, and basic concepts will stimulate further research.We are confident that the joint efforts ofthe authors and editors will contribute to a better understand- ing ofthe advances in the study ofthe challenging area ofsurface microbiol- ogy and will further stimulate progress in this field. It has been a pleasure to edit this book,primarily due to the stimulating cooperation ofthe contributors.We would like to express sincere thanks to all the staff members of Springer-Verlag, Heidelberg, especially, Drs. Dieter Czeschlik and Jutta Lindenborn for their help and active cooperation during the preparation ofthe book. New Delhi,India Ajit Varma Nedlands,Australia Lynette Abbott Marburg,Germany Dietrich Werner Tübingen,Germany Rüdiger Hampp July 2003 Contents 1 The State ofthe Art . . . . . . . . . . . . . . . . . . . . . . . 1 Ajit Varma,Lynette K.Abbott,Dietrich Werner andRüdiger Hampp Section A 2 Root Colonisation Following Seed Inoculation . . . . . . . 13 Thomas F.C.Chin-A-Woeng andBen J.J.Lugtenberg 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 2 Bacterial Root Colonisation . . . . . . . . . . . . . . . . . . 13 3 Analysis ofTomato Root Tip Colonisation After Seed Inoculation Using a Gnotobiotic Assay . . . . . . 14 3.1 Description ofthe Gnotobiotic System . . . . . . . . . . . . 14 3.2 Seed Disinfection . . . . . . . . . . . . . . . . . . . . . . . . 15 3.3 Growth and Preparation ofBacteria . . . . . . . . . . . . . . 16 3.4 Seed Inoculation . . . . . . . . . . . . . . . . . . . . . . . . 17 3.5 Analysis ofthe Tomato Root Tip . . . . . . . . . . . . . . . . 17 3.6 Confocal Laser Scanning Microscopy . . . . . . . . . . . . . 18 4 Genetic Tools for Studying Root Colonisation . . . . . . . . 18 4.1 Marking and Selecting Bacteria . . . . . . . . . . . . . . . . 18 4.2 Rhizosphere-Stable Plasmids . . . . . . . . . . . . . . . . . 21 4.3 Genetic and Metabolic Burdens . . . . . . . . . . . . . . . . 21 5 Behaviour ofRoot-Colonising PseudomonasBacteria in a Gnotobiotic System . . . . . . . 22 5.1 Colonisation Strategies ofBacteria . . . . . . . . . . . . . . 22 5.2 Competitive Colonisation Studies . . . . . . . . . . . . . . . 23 5.3 Monocots versus Dicots . . . . . . . . . . . . . . . . . . . . . 25 6 Influence ofAbiotic and Biotic Factors . . . . . . . . . . . . 25 VIII Contents 6.1 Abiotic Factors . . . . . . . . . . . . . . . . . . . . . . . . . 25 6.2 Biotic Factors . . . . . . . . . . . . . . . . . . . . . . . . . . 27 7 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 References and Selected Reading . . . . . . . . . . . . . . . . . . . . . 28 3 Methanogenic Microbial Communities Associated with Aquatic Plants . . . . . . . . . . . . . . . . . . . . . . . 35 RalfConrad 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 2 Role ofPlants in Emission ofCH to the Atmosphere . . . . 35 4 3 Role ofPhotosynthates and Plant Debris for CH Production 38 4 4 Methanogenic Microbial Communities on Plant Debris . . . 40 5 Methanogenic Microbial Communities on Roots . . . . . . . 42 6 Interaction ofMethanogens and Methanotrophs . . . . . . . 44 References and Selected Reading . . . . . . . . . . . . . . . . . . . . . 45 4 Role ofFunctional Groups ofMicroorganisms on the Rhizosphere Microcosm Dynamics . . . . . . . . . . 51 Galdino Andrade 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 2 General Aspects ofFunctional Groups ofSoil Microorganisms . . . . . . . . . . . . . . . . . . . . . 52 3 Carbon Cycle Functional Groups . . . . . . . . . . . . . . . 53 4 Functional Groups ofMicroorganisms ofthe Nitrogen Cycle 55 5 Functional Groups ofMicroorganisms ofthe Sulphur Cycle 57 6 Functional Groups ofMicroorganisms ofthe Phosphorus Cycle . . . . . . . . . . . . . . . . . . . . 59 7 Dynamics ofthe Rhizosphere Functional Groups ofMicroorganisms . . . . . . . . . . . . . . . . . . . . . . . 60 8 Relationship Among r and k Strategist Functional Groups . 61 9 Arbuscular Mycorrhizal Fungi Dynamics in the Rhizosphere . . . . . . . . . . . . . . . . . . . . . . . 61 10 Dynamics Among the Functional Micro-Organism Groups ofthe Carbon,Nitrogen,Phosphorus and Sulphur Cycles . . 65 References and Selected Reading . . . . . . . . . . . . . . . . . . . . . 68 Contents IX 5 Diversity and Functions ofSoil Microflora in Development ofPlants . . . . . . . . . . . . . . . . . . . . 71 Ramesh Chander Kuhad,David Manohar Kothamasi, K.K.Tripathi andAjay Singh 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . 71 2 Functional Diversity ofSoil Microflora . . . . . . . . . . . . 72 3 Role ofSoil Microflora in Plant Development . . . . . . . . 76 3.1 Mycorrhiza . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76 3.2 Actinorhiza . . . . . . . . . . . . . . . . . . . . . . . . . . . 80 3.3 Plant Growth-Promoting Rhizobacteria . . . . . . . . . . . 82 3.4 Phosphate-Solubilizing Microorganisms . . . . . . . . . . . 84 3.5 Lignocellulolytic Microorganisms . . . . . . . . . . . . . . . 85 4 Plant Growth Promoting Substances Produced by Soil Microbes . . . . . . . . . . . . . . . . . . . . . . . . . 88 5 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . 90 References and Selected Reading . . . . . . . . . . . . . . . . . . . . . 91 6 Signalling in the Rhizobia–Legumes Symbiosis . . . . . . . 99 Dietrich Werner 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . 99 2 The Signals from the Host Plants . . . . . . . . . . . . . . . 101 2.1 Phenylpropanoids:Simple Phenolics,Flavonoids and Isoflavonoids . . . . . . . . . . . . . . . . . . . . . . . . 102 2.2 Metabolization ofFlavonoids and Isoflavonoids . . . . . . . 104 2.3 Vitamins as Growth Factors and Signal Molecules . . . . . . 106 3 Signals from the Microsymbionts . . . . . . . . . . . . . . . 107 3.1 Nod Factors . . . . . . . . . . . . . . . . . . . . . . . . . . . 107 3.2 Cyclic Glucans . . . . . . . . . . . . . . . . . . . . . . . . . . 109 3.3 Lipopolysaccharides . . . . . . . . . . . . . . . . . . . . . . 110 3.4 Exopolysaccharides . . . . . . . . . . . . . . . . . . . . . . . 110 4 Signal Perception and Molecular Biology ofNodule Initiation . . . . . . . . . . . . . . . . . . . . . . . 111 References and Selected Reading . . . . . . . . . . . . . . . . . . . . . 114 X Contents Section B 7 The Functional Groups ofMicro-organisms Used as Bio-indicator on Soil Disturbance Caused by Biotech Products such as Bacillus thuringiensis and Bt Transgenic Plants . . . . . . . . . . . . . . . . . . . . 121 Galdino Andrade 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . 121 2 General Aspects ofBacillus thuringiensis . . . . . . . . . . . 122 3 Survival in the Soil . . . . . . . . . . . . . . . . . . . . . . . 123 4 History ofBacillus thuringiensis-Transgenic Plants . . . . . 124 5 Persistence ofthe Protein Crystal in the Soil . . . . . . . . . 125 6 Effect ofBacillus thuringiensisand Its Bio-insecticide Protein on Functional Soil Microorganism Assemblage . . . 126 References and Selected Reading . . . . . . . . . . . . . . . . . . . . . 130 8 The Use ofACC Deaminase-Containing Plant Growth-Promoting Bacteria to Protect Plants Against the Deleterious Effects ofEthylene . . . . . . . . . 133 Bernard R.Glick andDonna M.Penrose 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . 133 2 Ethylene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134 3 ACC Deaminase . . . . . . . . . . . . . . . . . . . . . . . . . 135 3.1 Treatment ofPlants with ACC Deaminase Containing Bacteria . . . . . . . . . . . . . . . . . . . . . . . 137 4 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . 140 References and Selected Reading . . . . . . . . . . . . . . . . . . . . . 141 9 Interactions Between Epiphyllic Microorganisms and LeafCuticles . . . . . . . . . . . . . . . . . . . . . . . . 145 Lukas Schreiber,Ursula Krimm andDaniel Knoll 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . 145 2 Physical and Chemical Parameters ofthe Phyllosphere . . . 147 3 LeafSurface Colonisation and Species Composition . . . . . 149 4 Alteration ofLeafSurface Wetting . . . . . . . . . . . . . . . 150 5 Interaction ofBacteria with Isolated Plant Cuticles . . . . . 152 6 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . 153 References and Selected Reading . . . . . . . . . . . . . . . . . . . . . 154 Contents XI 10 Developmental Interactions Between Clavicipitaleans and Their Host Plants . . . . . . . . . . . . . . . . . . . . . 157 James F.White Jr.,Faith Belanger,Raymond Sullivan, Elizabeth Lewis,Melinda Moy,William Meyer andCharles W.Bacon 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . 157 2 Endophyte/Epibiont Niche . . . . . . . . . . . . . . . . . . . 157 3 Coevolution ofClavicipitalean Fungi with Grass Hosts . . . 158 4 The Jump from Insects to Plants . . . . . . . . . . . . . . . . 158 4.1 Trans-Kingdom Jump . . . . . . . . . . . . . . . . . . . . . . 158 4.2 Intermediate Stages in the Transition to Plants . . . . . . . . 158 4.3 Parasitism ofGrass Meristematic Tissues . . . . . . . . . . . 160 5 Developmental Differentiation ofEndophytic and Epiphyllous Mycelium . . . . . . . . . . . . . . . . . . . 160 5.1 Plant Cell Wall Alteration . . . . . . . . . . . . . . . . . . . . 160 5.2 Endophytic Mycelial Growth . . . . . . . . . . . . . . . . . . 160 5.3 Control ofEndophytic Mycelial Development . . . . . . . . 163 5.4 Epiphyllous Mycelial Development . . . . . . . . . . . . . . 163 5.5 Expression ofFungal Secreted Hydrolytic Enzymes in Infected Plants . . . . . . . . . . . . . . . . . . . 164 6 Modifications ofPlant Tissues for Nutrient Acquisition . . . 165 6.1 Development ofthe Stroma in Epichloë . . . . . . . . . . . . 165 6.2 Stroma Development in Myriogenospora . . . . . . . . . . . 166 6.3 Mechanisms for Modifying Plant Tissues . . . . . . . . . . . 168 6.4 Evaporative-Flow Mechanism for Nutrient Acquisition . . . 169 6.5 The Cytokinin Induction Hypothesis . . . . . . . . . . . . . 169 7 Evolution ofAsexual Derivatives ofEpichloë . . . . . . . . . 171 7.1 Reproduction and Loss ofSexual Reproduction . . . . . . . 171 7.2 The Hypotheses . . . . . . . . . . . . . . . . . . . . . . . . . 172 7.3 The Process ofStroma Development and its Loss . . . . . . 173 7.4 The Shift from Pathogen to Mutualist . . . . . . . . . . . . . 174 8 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . 174 References and Selected Reading . . . . . . . . . . . . . . . . . . . . . 174 11 Interactions ofMicrobes with Genetically Modified Plants . 179 Michael Kaldorf,Chi Zhang,Uwe Nehls, Rüdiger Hampp andFrançois Buscot 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . 179 2 Changes in Microbial Communities Induced by Genetically Modified Plants . . . . . . . . . . . . . . . . . 181
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