Intestinal Microorganisms of Termites and Other Invertebrates

492 Pages·2006·10.73 MB·English

by  Helmut König

Soil Biology
6
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
Helmut König
Ajit Varma (Eds.)
Intestinal Microorganisms
of Termites and Other
Invertebrates
With86Figures,7inColor
123
Prof.Dr.HelmutKönig
JohannesGutenbergUniversity
InstituteofMicrobiologyandWineResearch
55099Mainz
Germany
e-mail:[email protected]
Prof.Dr.AjitVarma
Director
AmitySchoolofHerbalandMicrobialStudies
Sector125
NewSuperExpressHighway
Noida,201303,UP
India
e-mail:[email protected]
LibraryofCongressControlNumber:2005929881
ISSN1613-3382
ISBN-103-540-28180-0SpringerBerlinHeidelbergNewYork
ISBN-13978-3-540-28180-1SpringerBerlinHeidelbergNewYork
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Foreword
Insects and other invertebrates ranging from nematodes to earthworms,
isopods and millipeds usually host many microorganisms in both their
intestinal tracts and tissues. Outside of the gut these organisms are usu-
allyfoundinthehemolymphorarerestrictedtospecializedbacteriocytes
(mycetocytes)andmycetoms.Manymicroorganismshavebeencharacter-
ized morphologically (for example, compilation by Edward A. Steinhaus
1967) and their transfer to other developmental stages has been studied
thoroughly(PaulBuchner1953).Inthegutthisintra-andextracellularmi-
crobialdiversityisoftenreflectedbyculturableformsbutduetomolecular
techniquesmoreandmorespeciescanbeidentifiedtaxonomically.Italso
becameevidentthatarthropoddigestivetracts–withtheiroutpocketings
suchasforegutcaeca,ormalpighiantubulesandtheirunidirectionalbulk
flowofingestedfood–representprimaryhabitatsformicroorganismsin-
cluding bacteria, fungi, yeasts and protozoa. These organisms may occur
intraepithelially,adheretothegutwallorarefreefloatinginthelumenof
thedigestivetract.Numberandcompositionofmicroorganismscanvary
considerably, depending on longitudinal gut position, temperature, pH,
anaerobicoraerobicconditionsandvariousnaturalorlaboratorydiets.
Thesedigestivetractsarecharacterizedbyanexcesssupplyofmoreor
lessdegradablefood,controlledwateractivitiesandoftenratherstablepH
andtemperatureconditions.Ontheotherhandgutsmayrepresentunstable
environments because food composition may change in time and cuticle
liningsoffore-andhindgutsarelostwitheachmolt.Thisreestablishingof
microbiotaisespeciallyinterestinginholometabolousinsectswherelarvae
andadultsarefoundindifferenthabitats.
Theinteractionbetweenmicroorganismsandtheirhostandthehostguts
respectivelyissofarunknown.Theymaypassthroughthegutwithfood,
staythereforalongerperiodoftime(ascomparedwiththetransportation
of food) or represent true symbionts. In addition intracellular symbionts
fromoutsidethegutmustbetransferredthroughthedigestivetractatsome
pointinordertosettlewithinthebodycavity.The“symbiosis”-conceptcan
beseenfromabroadornarrowangle.Todaysymbiosishasstrongconnota-
tionsofstrictmutualismandcompletelyexcludesparasitism(assymbiosis
wasdefinedin1891byAntomdeBary(1831–1888)).PaulBuchner(1886–
VI Foreword
1978),whodedicatedmostofhislifetothestudyofendosymbiosis,defined
it ascohabitationbetween two partnersofdifferent species, where one is
takenupinthebodyoftheother,usuallymorehighlyorganised,partner.
Thebiologicalroleofthesemicroorganismsmayvaryconsiderably:gut
microorganisms of exotic hosts such as larvae of the oil fly Helaeomyia
petroleimightbeinterestingduetotheirpronouncedsolventtoleranceor
maybeafuturesourceofindustriallyusefulsolvent-tolerantenzymes.In
additionthesemicrobiotamaydetoxifyallelochemicalsormayberespon-
sibleforthebiosynthesisofessentialcompoundssuchasvitamins,sterols,
ornitrogen-containingconstituents.Generallythesemicroorganismsmay
be the source of novel metabolic capabilities such as defense substances
andantibioticsormayfixateatmosphericnitrogen.Ithasbeenknownfor
manyyearsthatenzymesinthegutwhichoriginatefromingestedfungal
tissueorgutmicrobesmaymediatecellulolysis.Alargenumberofinsects
andinvertebrateswhichusuallyutilizeinternalorexternalmicrobeshave
evolvedbeneficialassociationswiththesemicroorganisms.
TheeditorsHelmutKönigandAjitVarmadeservecreditforcompiling
articlesontheseinterestingphenomena.WithIntestinalMicroorganismsof
TermitesandOtherInvertebratesthey–togetherwiththeauthors–present
a fascinating and stimulating collection of articles focusing on intestinal
microorganismsofsoilinvertebratesandespeciallyoftermites,whichhave
beenstudiedextensively.Finally,thisbookintheseriesSoilBiologyishighly
innovativeincoveringbothmolecularandmicromethods.
IamsurethattherecentlyfoundedworkinggroupoftheGermanSociety
forGeneralandAppliedEntomology(DGaaE)entitled“Microbiologyand
Arthropods” together with this promising and comprehensive book will
trulyinspirethisfascinatingareaofinterdisciplinaryresearchandIthank
botheditorsandSpringerforrealizingthisproject.
Bayreuth,July2005 KonradDettner,
President
oftheGermanSociety
forGeneralandApplied
Entomology(DGaaE)
Preface
The soil is an important natural habitat for a large number of taxonomic
groups such as bacteria, archaea, protists, fungi, algae as well as plants
andanimals,whichhaveadaptedtotheenvironmentalconditionsinsoil.
It represents not only a steady state, but rather a constantly changing
environment, which is influenced by natural environmental factors and
humanactivities.
Soil biology describes the systematics, complex activities and interac-
tions of the soil inhabitants. A special discipline, the soil microbiology,
characterizesthemicrobialcommunityinthesubstratum.Itplaysamajor
role in the degradation and recycling of organic material. Microbes are
involvedinthefirststepofthesoilfoodweb,soilfertility,degradationof
xenobioticsaswellasinplantpathology.Inparticular,thedegradationof
lignocellulose, the most important renewable natural material, is mainly
adomainofmicroorganisms.Itisthereforeimperativetostudyecological
andagriculturalaspectsofsoilmicrobes.Inthepast,attentionwasmainly
directed towards the free-living or particle-bound microorganisms, and
theroleofintestinalmicrobesoccurringingutsystemsofsoilanimalshas
beenneglected.
The primary decomposers of organic soil litter are the lumbricids, the
diplopods,theisopods,anddipteranlarvaeaswellastermitesinsubtrop-
ical and tropical regions. Only a small percent of the soil microbes have
been screened axenically. The same is true for the intestinal microbes of
soil invertebrates. However, with the aid of molecular methods (e.g. total
16SrDNAsequenceanalysis),itwaspossibletogetaroughestimateofthe
totalpopulationinagivenenvironment.Inthelastfewyears,theintesti-
nal microbial communities (microbiota) of some soil invertebrates such
as collembola, earthworms, nematodes, isopods, millipedes and termites
have been studied in more detail. The most intensively studied group is
thatofthetermitesbecauseoftheirinterestingmicrofloraandglobalrole
inthedecompositionoforganicmaterial.
The intestinal microbiota has only been investigated from a few soil
invertebrates.Forthefirsttime,theauthorshighlightthisaspectandgive
anoverviewontheintestinalsoilcommunityinordertounderlinetheirrole
inthesoilfoodweb.Thegoalofthisbookistobridgeagapandtoaddanew
VIII Preface
mosaicstonetoourknowledgeofsoilbiology.Thebookisdividedintothree
sections.Thefirstdescribestheintestinalmicrobiotaoftheinvestigatedsoil
invertebrates,whiletheseconddealswithtermites,thebestinvestigatedsoil
invertebrateswithrespecttotheirgutmicrobiota.Thethirdpartpresents
noveltechniquesintroducedinecologicalmicrobiology,whichhavebeen
successfullyappliedtostudiesoftheintestinalmicrobiota.Thebookcovers
severalnovelfacetsandpresentsinsightsintotheintestinalmicrobiotaof
soil invertebrates. The text is mainly directed towards graduate students
and professional scientists with a general interest in the soil or intestinal
microbiotaorinecologicalmicrobiologyingeneral.
Wearegratefultoallauthorsforprovidingtheirexpertisesandcontri-
butions,makingthisspecialeditionpossible.We alsoexpressourthanks
toSpringer-Verlagforacceptingthisunconventionalandlessinvestigated,
butecologicallyimportant,subjectforpublicationinthisprestigiousbook.
WeappreciatetheeffortsofDr.DieterCzeschlikandDr.JuttaLindenborn
fromSpringerLifeSciencesEditorial,whomanagedthisspecialbook.
MainzandNoida,April2005 HelmutKönigandAjitVarma
Contents
Part I Soil Invertebrates
1 BiologyofSoilInvertebrates 3
GerhardEisenbeis
1.1 Introduction.................................................................. 3
1.2 TheMicrofauna.............................................................. 8
1.2.1 Protozoa............................................................... 8
1.2.2 Nematoda–Roundworms,Eelworms....................... 10
1.3 TheMesofauna............................................................... 12
1.3.1 Pseudoscorpionida–FalseScorpions,BookScorpions 13
1.3.2 Acari–Mites......................................................... 14
1.3.3 Symphyla............................................................. 17
1.3.4 Pauropoda............................................................ 19
1.3.5 Collembola–Springtails......................................... 19
1.3.6 Protura................................................................ 23
1.3.7 Diplura–DoubleTails............................................ 25
1.4 TheMacrofauna ............................................................. 26
1.4.1 Araneida–Spiders................................................. 27
1.4.2 Opiliones–Harvestmen......................................... 29
1.4.3 TerrestrialIsopoda(Oniscoidea)–Woodlice............. 30
1.4.4 Chilopoda–Centipedes.......................................... 33
1.4.5 Diplopoda-Millipedes........................................... 35
1.4.6 Enchytraeidae–Whiteworms,Potworms.................. 37
1.4.7 Oligochaeta:Lumbricidae–Earthworms .................. 39
1.4.8 TerrestrialGastropoda–SlugsandSnails.................. 41
1.4.9 Insecta–PterygoteInsects(ShortCommentsAbout
theRoleofSelectedGroupsofHigherInsects)........... 42
1.5 Conclusions................................................................... 43
References............................................................................. 47
2 InteractionsBetweenBacteriaandNematodes 55
LeoEberl,DavidJ.Clarke
2.1 Introduction.................................................................. 55