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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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Soils would not exist without the complex and heterogeneous activities of microorganisms. For the third volume of Soil Biology, an international board of renowned scientists shed light on the significant role of these organisms. The following key topics are covered: Microorganisms in bioerosion, hum
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