Microbial Root Endophytes PDF
Preview Microbial Root Endophytes
Soil Biology 9 Series Editor: Ajit Varma Volumespublishedintheseries Volume1 A.Singh,O.P.Ward(Eds.) AppliedBioremediationandPhytoremediation 2004 Volume2 A.Singh,O.P.Ward(Eds.) BiodegradationandBioremediation 2004 Volume3 F.Buscot,A.Varma(Eds.) MicroorganismsinSoils:RolesinGenesisandFunctions 2005 Volume4 S.Declerck,D.-G.Strullu,J.A.Fortin(Eds.) InVitroCultureofMycorrhizas 2005 Volume5 R.Margesin,F.Schinner(Eds.) ManualforSoilAnalysis– MonitoringandAssessingSoilBioremediation 2005 Volume6 H.König,A.Varma(Eds.) IntestinalMicroorganismsofTermites andOtherInvertebrates 2006 Volume7 K.G.Mukerji,C.Manoharachary,J.Singh(Eds.) MicrobialActivityintheRhizosphere 2006 Volume8 P.Nannipieri,K.Smalla(Eds.) NucleicAcidsandProteinsinSoil 2006 Barbara J.E. Schulz Christine J.C. Boyle Thomas N. Sieber (Eds.) Microbial Root Endophytes With29Figures,4inColor 123 PDDr.BarbaraJ.E.Schulz Dr.ChristineJ.C.Boyle TechnicalUniversityofBraunschweig Augustastraße32 InstituteofMicrobiology 02826Görlitz Spielmannstraße7 Germany 38106Braunschweig e-mail:[email protected] Germany e-mail:[email protected] Dr.ThomasN.Sieber SwissFederalInstituteofTechnology DepartmentofEnvironmentalSciences InstituteofIntegrativeBiology ForestPathologyandDendrology 8092Zürich Switzerland e-mail:[email protected] LibraryofCongressControlNumber:2005938057 ISSN1613-3382 ISBN-103-540-33525-0SpringerBerlinHeidelbergNewYork ISBN-13978-3-540-33525-2SpringerBerlinHeidelbergNewYork This work is subject to copyright. All rights reserved, whether the whole or part of the materialisconcerned,specificallytherightsoftranslation,reprinting,reuseofillustrations, recitation,broadcasting,reproductiononmicrofilmorinanyotherway,andstorageindata banks.Duplicationofthispublicationorpartsthereofispermittedonlyundertheprovisions oftheGermanCopyrightLawofSeptember9,1965,initscurrentversion,andpermission forusemustalwaysbeobtainedfromSpringer.Violationsareliableforprosecutionunder theGermanCopyrightLaw. SpringerisapartofSpringerScience+BusinessMedia springer.com ©Springer-VerlagBerlinHeidelberg2006 PrintedinGermany Theuseofgeneraldescriptivenames,registerednames,trademarks,etc.inthispublication doesnotimply,evenintheabsenceofaspecificstatement,thatsuchnamesareexempt fromtherelevantprotectivelawsandregulationsandthereforefreeforgeneraluse. Editor:Dr.DieterCzeschlik,Heidelberg,Germany DeskEditor:Dr.JuttaLindenborn,Heidelberg,Germany Coverdesign:design&production,Heidelberg,Germany Typesettingandproduction:LE-TEXJelonek,Schmidt&VöcklerGbR,Leipzig,Germany 31/3150-YL-543210-Printedonacid-freepaper Preface Healthy plant roots are not only colonized by mycorrhizal fungi and rhi- zobial bacteria, but also by a myriad of other microorganisms, including endophyticbacteriaandfungi.Comparativelylittleisknownaboutthese endophytic microorganisms, which do not cause apparent disease, but colonizeroottissuesinter-and/orintracellulary.Althoughtherehadbeen previous research studying both bacterial and fungal endophytes, it was inthemid-1980sthatnumerousinvestigatorsbeganstudyingthesegroups of microorganisms more intensively. Initially, most work on endophytes centered on the diversity of isolates and correlations with ecological fac- tors. Recently it has become clear that some of these interactions with endophyticbacteriaandfungicanbelatentlypathogenicand/ormutualis- tic.Inmutualisticinteractions,theendophytemayimprovegrowthofthe host,conveystresstolerance,inducesystemicresistance,orsupplythehost withnutrients.Ontheotherhand,mostendophytesarealsoabletogrow saprotrophically,e.g.,fromsurface-sterilizedtissuesonmediacontaining deadorganicsubstrates.Thus,ithasbecomeobviousthatendophyteshave multiple life history strategies and that these can be extremely plastic, as willbecomecleartothereadersofthesubsequent19chapters. Thisbookisthefirsttodealwithbacterialandfungalrootendophytes, theirdiversity,lifehistorystrategies,interactions,applicationsinagricul- tureandforestry,andalsowithmethodsforisolation,cultivation,andboth conventionalandmolecularmethodsforidentificationanddetection.The first chapter deals with the question: What are endophytes? However, it alsointroducesthereadertothesubjectstreatedinthesubsequentchap- ters. We hope that readers will not only find this book informative, but will also be provoked to further study these fascinating interactions, and inparticulartobetterunderstandthemechanismsregulatingthem.Itwill becomeapparentthatwearestillfarfromunderstandingthefactorsthat determine whether a plant-microbial interaction remains asymptomatic, leadstodisease,orismutualistic. VI Preface Wewouldliketothankourcolleaguesfortheircontributionsandtheir worktomakethisbookasuccessfulunity,toJuttaLindenbornofSpringer forherfriendlyhelpandadvice,andtoAjitVarmafortheinvitationtoedit abookinthisseries. Braunschweig, BarbaraSchulz GörlitzandZürich, ChristineBoyle June2006 andThomasSieber Contents 1 WhatareEndophytes? 1 BarbaraSchulz,ChristineBoyle 1.1 IntroductionandDefinitions............................................ 1 1.2 Colonisation.................................................................. 2 1.3 AssemblagesandAdaptation............................................ 4 1.4 LifeHistoryStrategies ..................................................... 6 1.5 BalancedAntagonism...................................................... 7 1.6 Conclusions................................................................... 9 References............................................................................. 10 Part I Endophytic Bacteria 2 SpectrumandPopulationDynamics ofBacterialRootEndophytes 15 JohannesHallmann,GabrieleBerg 2.1 Introduction.................................................................. 15 2.2 PopulationDensity ......................................................... 15 2.3 BacterialSpectrum ......................................................... 16 2.4 BacterialDiversity.......................................................... 21 2.5 FactorsInfluencingColonisation....................................... 21 2.5.1 Methodology....................................................... 21 2.5.2 Geography .......................................................... 22 2.5.3 PlantSpecies ....................................................... 22 2.5.4 PlantGenotype.................................................... 23 2.6 Interactions................................................................... 24 2.6.1 PlantPathogens................................................... 24 2.6.2 PlantSymbionts................................................... 25 2.6.3 PlantDefenceMechanisms.................................... 25 2.6.4 AgriculturalPractices........................................... 25 2.7 PotentialHumanPathogensAmongRootEndophytes.......... 26 2.8 Conclusions................................................................... 27 References............................................................................. 28 VIII Contents 3 BacterialEndophytesasElicitorsofInducedSystemicResistance 33 JosephW.Kloepper,Choong-MinRyu 3.1 IntroductionandTerminology ......................................... 33 3.2 ScopeofEndophytesthatElicitInducedResistance andPathosystemsAffected............................................... 34 3.3 InternalColonizationofEndophytes thatElicitInducedResistance........................................... 39 3.4 PlantResponsestoEndophyticElicitors............................. 41 3.5 ImplementationinProductionAgriculture: TwoCaseStudies............................................................ 44 3.6 Conclusions................................................................... 49 References............................................................................. 50 4 ControlofPlantPathogenicFungiwithBacterialEndophytes 53 GabrieleBerg,JohannesHallmann 4.1 Introduction.................................................................. 53 4.2 SpectrumofIndigenousEndophyticBacteria withAntagonisticPotentialTowardsFungalPlantPathogens 54 4.3 ModeofActionofAntagonisticBacteria............................ 58 4.3.1 Antibiosis ........................................................... 58 4.3.2 Competition........................................................ 59 4.3.3 Lysis................................................................... 59 4.3.4 InductionofPlantDefenceMechanisms.................. 60 4.3.5 PlantGrowth....................................................... 60 4.4 ControlPotentialofEndophyticBacteria............................ 60 4.5 EnhancingBiocontrolEfficiency....................................... 61 4.6 Conclusions................................................................... 65 References............................................................................. 66 5 RoleofProteinsSecretedbyRhizobiainSymbiotic InteractionswithLeguminousRoots 71 MagedM.Saad,WilliamJ.Broughton,WilliamJ.Deakin 5.1 Introduction.................................................................. 71 5.2 BacterialProteinSecretionSystems................................... 73 5.2.1 TypeISecretionSystems....................................... 73 5.2.2 TypeIISecretionSystems...................................... 77 5.2.3 TypeIIISecretionSystems..................................... 77 5.2.4 TypeIVSecretionSystems..................................... 82 5.3 Conclusions................................................................... 83 References............................................................................. 83 Contents IX 6 ResearchonEndophyticBacteria:RecentAdvances withForestTrees 89 RichaAnand,LesliePaul,ChrisChanway 6.1 Introduction.................................................................. 89 6.2 BacterialEndophytesofForestTrees.................................. 91 6.3 EndophyticBacteriaofConifers........................................ 92 6.4 ModesandSitesofEntry.................................................. 95 6.5 MechanismsofPlantGrowthPromotion............................ 97 6.6 FutureWork................................................................... 102 References............................................................................. 103 Part II Endophytic Fungi 7 BiodiversityofFungalRoot-EndophyteCommunities andPopulations,inParticularoftheDarkSeptateEndophyte Phialocephalafortiniis.l. 107 ThomasN.Sieber,ChristophR.Grünig 7.1 Introduction.................................................................. 107 7.2 SpeciesDiversityofRootEndophyteCommunities.............. 108 7.2.1 GeographyandClimate......................................... 109 7.2.2 Soil .................................................................... 114 7.2.3 MultitrophicInteractions ...................................... 115 7.2.4 NaturalandAnthropogenicDisturbances................ 117 7.3 DarkSeptateEndophytes................................................. 119 7.3.1 History............................................................... 119 7.3.2 Biodiversity......................................................... 119 7.3.3 DiversityofPhialocephalafortinii........................... 121 7.4 Conclusions................................................................... 125 References............................................................................. 126 8 EndophyticRootColonizationbyFusariumSpecies: Histology,PlantInteractions,andToxicity 133 CharlesW.Bacon,IdaE.Yates 8.1 Introduction.................................................................. 133 8.2 PlantandFungusInteractions.......................................... 134 8.2.1 HemibiotrophicCharacteristics.............................. 138 8.2.2 Histology............................................................ 139 8.2.3 Mycotoxins.......................................................... 143 8.2.4 MycotoxinsandHostRelationships......................... 144 8.2.5 PhysiologicalInteractionsandDefenseMetabolites... 145 8.3 Summary....................................................................... 146 References............................................................................. 147 X Contents 9 MicrobialEndophytesofOrchidRoots 153 PaulBayman,J.TupacOtero 9.1 Introduction.................................................................. 153 9.2 HabitsandTypesofOrchidRoots ..................................... 153 9.3 BacteriaasEpiphytesandEndophytesofOrchidRoots ........ 154 9.4 OrchidEndophytesorOrchidMycorrhizalFungi?............... 155 9.5 ProblemswiththeTaxonomyofOrchidEndophyticFungi.... 157 9.6 HostSpecificityofOrchidEndophytes............................... 158 9.7 EndophyticFungiinRootsofTerrestrial, PhotosyntheticOrchids................................................... 158 9.8 EndophyticFungiinRootsofMyco-HeterotrophicOrchids.. 167 9.9 EndophyticFungiinRootsofEpiphytic andLithophyticOrchids.................................................. 169 9.10 EndophyticFungiinEpiphyticOrchidRoots: ImportancetoPlantHosts................................................ 171 9.11 Conclusions................................................................... 172 References............................................................................. 173 10 FungalEndophytesinSubmergedRoots 179 FelixBärlocher 10.1 Introduction.................................................................. 179 10.2 AquaticHyphomycetes.................................................... 180 10.3 FungiinSubmergedRoots............................................... 181 10.4 ConclusionsandOutlook................................................. 186 References............................................................................. 188 11 NematophagousFungiasRootEndophytes 191 LuisV.Lopez-Llorca,Hans-BörjeJansson,JoséGasparMaciáVicente, JesúsSalinas 11.1 Introduction.................................................................. 191 11.2 NematophagousFungi..................................................... 191 11.2.1 NematodeParasites.............................................. 192 11.2.2 Mycoparasites...................................................... 195 11.2.3 RootEndophytes.................................................. 195 11.3 ConcludingRemarks....................................................... 202 References............................................................................. 203 12 MolecularDiversityandEcologicalRolesofMycorrhiza-Associated SterileFungalEndophytesinMediterraneanEcosystems 207 MariangelaGirlanda,SilviaPerotto,AnnaMariaLuppi 12.1 Introduction.................................................................. 207
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