Table Of ContentSoil 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
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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
Description: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