Soil Biology 3 Series Editor: Ajit Varma Franc¸ois Buscot • Ajit Varma (Eds.) Microorganisms in Soils: Roles in Genesis and Functions With71Figures 123 ProfessorDr.AjitVarma ProfessorDr.Franc¸oisBuscot JawaharlalNehruUniversity UniversitätLeipzig SchoolLifeScience InstitutfürBotanik Mol.GeneticsLab. Abt.TerrestrischeÖkologie NewMehrauliRoad Johannisallee21 110067NewDelhi 04103Leipzig India Germany e-mail:[email protected] and AmityInstitute ofHerbalandMicrobialStudies Sector125 NewSuperExpressHighway Noida India e-mail:[email protected] LibraryofCongressControlNumber:2004111759 ISSN1613-3382 ISBN3-540-22220-0SpringerBerlinHeidelbergNewYork 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 springeronline.com ©Springer-VerlagBerlinHeidelberg2005 PrintedinGermany Theuseofgeneraldescriptivenames,registerednames,trademarks,etc.inthispublication doesnotimply,evenintheabsenceofaspecificstatement,thatsuchnamesareexempt fromtherelevantprotectivelawsandregulationsandthereforefreeforgeneraluse. Coverdesign:design&production,Heidelberg,Germany Typesettingandproduction:LE-TEXJelonek,Schmidt&VöcklerGbR,Leipzig,Germany 31/3150-YL-543210-Printedonacid-freepaper Preface The inspiration and concept for preparing this volume were conceived while both of us were sipping a cup of Columbian coffee in the Goethe Gallery, Jena (Germany). The idea developed that soils are dynamic bio- logicalsystemsandcertainlynotastaticsubstratethatsupportsthelifeof microbes,plantsandanimals.Microorganismsplayavitalroleincreating thisuniverseandmaintaininglifeinit.Dealingwiththeinterrelationships oforganismsandtherelationshipsbetweenorganismsandtheirenviron- ment was formerly more or less confined to a small group of specialists within the scientific community. This has changed in the current sce- nario. Growing environmental problems have created a public awareness of the ecological disturbances and dangers related to excessive industrial expansionand theway of life in“disposablesocieties”. As a consequence of the perceived importance of ecology, research in this field has devel- opedrapidly.Asoneofthethreeenvironmentalmediabesideswaterand air, soils have now become a central concern for a broad range of scien- tists. In the golden era of microbiology, the study of soil organisms soon becameanareaofinteresttoalargenumberofearlybacteriologists,andthe pioneeringinvestigationsofWinogradsky,Omeliansky,andBeijerinckstill standasmajorcontributionstoourknowledgeofthebacterialpopulation. At the same time, it became apparent to soil scientists that the surface crustoftheearthisnotmerelyastaticphysiochemicalmatrixuponwhich green plants grow, but also a biological system in a continuous dynamic equilibrium. In the realm of pure science, information on the ecology, function, and biochemistry of microflora has grown considerably so that aclearpictureofsoilbiologyisbeginningtoemerge. Theinnumerabledevelopmentsinrecentyearsmakeacompletereview impossiblewithinthescopeofasinglevolume.Someofthemoredetailed points have been omitted for brevity, yet, where conflicts still exist, con- trastingviewpointsarepresented. Timemaychangetheseviews,butitis in the very nature of science to be in a continual state of flux and for the errors of one generation to be amended by the next. Soil microbiologyis not a pure discipline. Its parentage may be traced through bacteriology, mycology, and soil science; biochemistry and plant pathology have also madetheirmark,especiallyinrecentyears. VI Preface Intheframeworkofagriculture,themicrofloraisofsignificanceforman’s ability to feed himself. For the microbial inhabitant, the soil functions as auniqueecosystemtowhichtheorganismmustadaptandfromwhichit mustobtainsustenance.However,inthefinalanalysis,microbiologistscan find definitive answers as to how these processes are brought about only throughbiochemicalinquiries. We have attempted to bring together the major aspects of rhizosphere researchandprinciplesofrhizosphereecologyforthebenefitofdeveloping youngscientistsandtechnologies,aswellasfortheestablishedprofessional researcherandteacher.Aprimeobjectiveandhopeisthatthisvolumemight generateideasthatwillbringforthnewapproachesandmethodologylead- ing to further advances in our understanding of rhizosphere interactions and their implications for agriculture and forestry. Nevertheless, even if therhizosphereisthecompartmentfromwhichplantsacquiretheirwater and nutrients and a hot spot of microbial and animal activity, this com- partmentcanonlybeunderstoodinthecontextofwholesoilfunctioning, fromsoilgenesistothenutrientcycling,andincludingtheexchangeswith waterandatmosphere.Theseaspectsthereforeoccupyalargepartofthe volume. Referencesareofgreatvaluenotonlytotheresearchworker,buttothe advancedstudentaswell.Theblindacceptanceofsecondarysourceswhen primary material is readily accessible is not the hallmark of the serious student.Whereavailable,reviewsareincludedinthereferencelistsofeach chaptersothatthefinerpointsofeachtopicmaybesoughtout.Pertinent originalcitationsarelikewiseincludedsincethesepermitthestudentand researcheraliketoexaminetheoriginalsource,observethetechniquesuti- lized,anddrawtheirownconclusions.However,amereliteraturereviewis notintended,sincemuchgoodworkhasnotbeencited.Wehavedeliber- atelydrawnuponsomeoldresearchinformation,largelyforthebenefitof advancedstudentsandyoungscientists,toshowwhereresearchhascome fromandwhereitmaybegoing.Indoingthis,webelievewehaverevealed manygapsinourknowledgewhichareyettobefilled.Emphasisisgiven to the more recent papers, but certain classical works are also included, particularly where the studies have been of such a nature as to define a unique approach. It may also be of value to students majoring in other fields,suchassoilscience,geology,hydrology,plantecology,zooecology, phytopathology,agronomy,forestry, or the environmental, cropsciences, naturalsciencemanagement,agriculturalengineering,biologicalsciences, animalsciences,andlifesciences. For meaningful contributions to be made in the future, the need for refinedtechnologyandamultidisciplinarypoolingofexpertisebysoilmi- crobiologists,phytopathologists,soilphysicistsandchemists,plantphysi- ologists,andzoologistsshouldbeclearlyevident. Preface VII Thepresentationisessentiallyarrangedintosixmainparts.TheIntro- duction or Part I outlines the definition of soils and dynamics to the mi- crobialdiversity.PartIIdealswithvariedfunctionsofthemicroorganisms andsoilgenesis.InPartIII,wehighlightthebiogeochemicalprocesses.The bioticinteractionsintermsofplant/microorganismsinvolvingsymbiosis aregiveninPartIV.Functionsofmicrobesinspecificsoilcompartments are discussed in Part V. Modern tools and techniques to understand soil biologyareelucidatedinPartVI. Itishopedthatthegroundworkwillbelaidhereinforafullerenquiry onthepartofthereaders.Ifthisgoalisachievedeveninpart,thevolume willhaveserveditspurpose. Molecularmicrobiologicalstudieshavefocusedourattentioninrecent times on the characterization of known as well as unknown microbial speciesimplicatedinsoiltransformationandplantgrowth. While assuming sole responsibility for any omissions or errors in the book,wearegratefultoallthoseunselfishindividualswhohavecontributed tothechapters.Finally,wewouldliketoaskthereadertomakeallowances for our lack of linguistic proficiency considering that English is not our mothertongue. Wearegratefultothemanypeoplewhohelpedbringthisvolumetolight. WewishtothankDr.DieterCzeschlikandDr.JuttaLindenborn,Springer- Verlag, Heidelberg for generous assistance and patience in finalizing the volume.Finally,specificthanksgotoourfamily,immediate,andextended, notforgettingthememoryofthosewhopassedaway,fortheirsupportor theirincentive in putting everything together. Ajit Varmain particularis very thankful to Dr. Ashok K. Chanhan, founder president, An Institute ofRitnandBalvedEducationFondation(Amity),NewDelhi,forthekind support and constant encouragement received. Special thanks are due to myPh.D.studentsMs.RinaKamariandMr.RamPrasadforcompilingthe subjectindex. Leipzig,Germany Fran¸coisBuscot NewDelhi,India AjitVarma June2004 Contents Part I Introduction 1 WhatAreSoils? 3 Fran¸coisBuscot 1 Introduction.................................................................. 3 2 SoilGenesis ................................................................... 4 2.1 RockWeatheringorDecay...................................... 4 2.2 ImportanceofSoilTexture...................................... 5 2.3 InputofOrganicMatterintoSoilsandAggregation.... 7 2.4 MigrationProcesses............................................... 8 3 BiogeochemicalProcessesinSoils..................................... 8 3.1 EnergyandCarbon................................................ 8 3.2 NitrogenandPhosphorus....................................... 10 4 BioticInteractionsInvolvingSoilMicroorganisms............... 11 4.1 CompetitionVersusFacilitation............................... 11 4.2 TheExampleofMycorrhizas................................... 12 5 IntegrativeConsiderationsonFunctionsofMicroorganisms inSpecificSoilCompartments.......................................... 13 5.1 ReleaseofTransgenicOrganismsasaTool toTraceEffectsofEcologicalDisruptions onSoilMicroorganisms.......................................... 13 5.2 SoilPollutionbyHeavyMetals asaMoreComplexDisruption................................ 14 5.3 UnderstandingComplexFunctionalDomains inSoilHabitats...................................................... 15 6 ConclusionorBacktoBiodiversityofSoilMicrobes............. 15 References............................................................................. 16 2 MicrobialDiversityinSoils 19 Bhoopander Giri, Pham Huong Giang, Rina Kumari, Ram Prasad, AjitVarma 1 Introduction.................................................................. 19 2 OriginofMicrobialDiversity............................................ 20 X Contents 2.1 OxygenRevolution ................................................ 21 2.2 OriginoftheFirstEukaryotes.................................. 22 3 TypesofSoilMicroorganisms........................................... 22 3.1 Eubacteria............................................................ 24 3.2 Archaebacteria...................................................... 29 3.3 Fungi................................................................... 31 3.4 Algae................................................................... 33 4 MicrobialDiversityandBiologicalSpheres......................... 33 4.1 TheDetritusphere ................................................. 34 4.2 TheDrilosphere.................................................... 34 4.3 ThePorosphere..................................................... 35 4.4 TheAggregatusphere............................................. 35 4.5 TheRhizosphere ................................................... 36 5 MicrobialDiversityandChemicalTransformation............... 37 5.1 NitrogenTransformation........................................ 38 5.2 PhosphorusTransformation.................................... 39 5.3 SulfurTransformation............................................ 41 5.4 IronTransformation.............................................. 42 6 MicrobialDiversityandBioticInteractions......................... 42 7 Conclusion .................................................................... 47 References............................................................................. 49 Part II Microorganisms and Soil Genesis 3 RoleofMicroorganismsinWearDownofRocksandMinerals 59 AnnaA.Gorbushina,W.E.Krumbein 1 RockWeatheringorRockWearDown? .............................. 59 2 CarbonDioxideandRockWearDown............................... 63 3 BalanceofCarbonDioxideSourcesandSinks..................... 68 4 RockWearDownasaPotentialCarbonDioxideSink........... 70 5 TheFractalDimensionofBiologicalRockWearDown......... 71 6 CalciumCarbonateandSilicateWearDown, DissolutionandPrecipitation WithSpecialReferencetoBiologicalRockDegradation........ 74 7 Conclusions................................................................... 79 References............................................................................. 80 4 HumificationandMineralizationinSoils 85 GeorgGuggenberger 1 DefinitionsandIntroduction............................................ 85 2 SoilOrganicMatterResources.......................................... 86 Contents XI 2.1 PlantCompounds.................................................. 87 2.2 MicrobialCompounds............................................ 89 2.3 BlackCarbon........................................................ 90 3 MineralizationandHumificationPathways......................... 91 3.1 FactorsAffectingDecompositionandMineralization.. 92 3.2 HumificationProcesses .......................................... 95 4 Conclusions................................................................... 102 References............................................................................. 104 5 ImportanceofMicroorganismsforSoilAggregation 107 Jean-LucChotte 1 Introduction.................................................................. 107 2 EvidenceoftheRoleofSoilMicroorganisms....................... 108 3 MicrobialMetabolitesResponsibleforSoilAggregation ....... 110 3.1 Polysaccharides..................................................... 110 3.2 Glomalin.............................................................. 111 3.3 Lipids .................................................................. 112 4 ManipulationofMicrobiallyMediatedProcesses toImproveSoilAggregation............................................. 113 4.1 TheRhizosphereMicrobialCommunity.................... 113 4.2 OrganicResidues................................................... 113 4.3 InoculationwithMicroorganisms............................ 114 5 Conclusion .................................................................... 115 References............................................................................. 115 Part III Microorganisms and Biogeochemical Processes in Soils 6 MicrobialEnergeticsinSoils 123 OliverDilly 1 Introduction.................................................................. 123 2 Soil,EnergyandMicroorganisms...................................... 124 3 MicrobialCommunities................................................... 127 4 MicrobialMetabolisminSoil............................................ 129 4.1 Catabolism........................................................... 129 4.2 Anabolism............................................................ 131 4.3 Soil OrganicC, MicrobialC and BiologicalActive C andInteractionswithN.......................................... 133 5 HolisticApproachestoEvaluateEnergeticStrategies ofSoilMicrobialCommunities ......................................... 133 6 Conclusions................................................................... 136 References............................................................................. 136 XII Contents 7 RoleofMicroorganismsinCarbonCyclinginSoils 139 EllenKandeler,MichaelStemmer,MartinH.Gerzabek 1 Introduction.................................................................. 139 2 CarbonSources.............................................................. 140 3 SpatialDistributionandProtectionofCarbonSources......... 142 4 SpatialDistributionofSoilMicroorganisms andTheirActivities......................................................... 143 5 MicroorganismsandEnzymesInvolvedinCCycling ........... 147 6 DynamicsofOrganicMatterDecomposition inAgroecosystems.......................................................... 148 7 SoilOrganicMatter,Below-GroundProcesses andClimateChange........................................................ 151 References............................................................................. 153 8 ContributionofBacteriatoInitialInputandCyclingofNitrogen inSoils 159 LaurentPhilippot,J.C.Germon 1 Introduction.................................................................. 159 2 NitrogenTransformationsintheSoil................................. 160 3 BacteriaInvolvedintheNitrogenCycle.............................. 162 3.1 Nitrogen-FixingBacteria........................................ 162 3.2 Nitrifiers.............................................................. 164 3.3 NitrateReducers,Denitrifiers andNitriteAmmonifiers......................................... 165 4 NitrogenFluxes.............................................................. 167 4.1 BiologicalNitrogenFixation.................................... 168 4.2 NitrogenMineralization......................................... 169 4.3 Nitrification.......................................................... 169 4.4 DissimilatoryNitrateReductiontoAmmonium......... 170 4.5 Denitrification...................................................... 171 References............................................................................. 172 9 InfluenceofMicroorganismsonPhosphorusBioavailability inSoils 177 AnnetteDeubel,WolfgangMerbach 1 Introduction.................................................................. 177 2 MicrobialEffectsonRhizodeposition................................ 177 3 MechanismsofMicrobialInfluence onPhosphorusAvailability............................................... 179 3.1 SolubilizationofCalciumPhosphates....................... 179 3.2 MobilizationofIron-andAluminum-Bound Phosphorus.......................................................... 181
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