(Endo)symbiotic Methanogenic Archaea PDF
Preview (Endo)symbiotic Methanogenic Archaea
Microbiology Monographs Volume 19 Series Editor: Alexander Steinbu¨chel Mu¨nster,Germany Microbiology Monographs Volumes published in the series InclusionsinProkaryotes UncultivatedMicroorganisms VolumeEditor:JessupM.Shively VolumeEditor:SlavaS.Epstein Vol.1,2006 Vol.10,2009 ComplexIntracellularStructures MicrobialMegaplasmids inProkaryotes VolumeEditor:EdwardSchwartz VolumeEditor:JessupM.Shively Vol.11,2009 Vol.2,2006 EndosymbiontsinParamecium MagnetoreceptionandMagnetosomes VolumeEditor:MasahiroFujishima inBacteria Vol.12,2009 VolumeEditor:DirkSchu¨ler Vol.3,2007 Alginates:BiologyandApplications VolumeEditor:BerndH.A.Rehm PredatoryProkaryotes–Biology,Ecology Vol.13,2009 andEvolution VolumeEditor:EdouardJurkevitch Vol.4,2007 PlasticsfromBacteria:NaturalFunctions andApplications VolumeEditor:Guo-QiangChen AminoAcidBiosynthesis–Pathways, Vol.14,2010 RegulationandMetabolicEngineering VolumeEditor:VolkerF.Wendisch Vol.5,2007 Amino-AcidHomopolymersOccurring inNature MolecularMicrobiologyofHeavyMetals VolumeEditor:YoshimitsuHamano VolumeEditors:DietrichH.Nies Vol.15,2010 andSimonSilver Vol.6,2007 BiologyofRhodococcus VolumeEditor: He´ctorAlvarez Vol.16,2010 MicrobialLinearPlasmids VolumeEditors:FriedhelmMeinhardt StructuresandOrganellesinPathogenic andRolandKlassen Protists Vol.7,2007 VolumeEditor:W.deSouza Vol.17,2010 ProkaryoticSymbiontsinPlants VolumeEditor:KatharinaPawlowski PlantGrowthandHealthPromotingBacteria Vol.8,2009 VolumeEditor:DineshK.Maheshwari Vol.18,2010 HydrogenosomesandMitosomes: (Endo)symbioticMethanogenicArchaea MitochondriaofAnaerobicEukaryotes VolumeEditor:JanTachezy VolumeEditor:JohannesH.P.Hackstein Vol.9,2008 Vol.19,2010 Johannes H.P. Hackstein Editor (Endo)symbiotic Methanogenic Archaea Editor Dr.JohannesH.P.Hackstein IWWR FacultyofScience RadboudUniversityNijmegen Heyendaalseweg135 6525AJNijmegen TheNetherlands [email protected] SeriesEditor ProfessorDr.AlexanderSteinbu¨chel Institutfu¨rMolekulareMikrobiologieundBiotechnology Westfa¨lischeWilhelms-Universita¨t Corrensstr.3 48149Mu¨nster Germany [email protected] ISSN1862-5576 e-ISSN1862-5584 ISBN978-3-642-13614-6 e-ISBN978-3-642-13615-3 DOI10.1007/978-3-642-13615-3 SpringerHeidelbergDordrechtLondonNewYork LibraryofCongressControlNumber:2010935950 #Springer-VerlagBerlinHeidelberg2010 Thisworkissubjecttocopyright.Allrightsarereserved,whetherthewholeorpartofthematerialis concerned,specificallytherightsoftranslation,reprinting,reuseofillustrations,recitation,broadcasting, reproductiononmicrofilmorinanyotherway,andstorageindatabanks.Duplicationofthispublication orpartsthereofispermittedonlyundertheprovisionsoftheGermanCopyrightLawofSeptember9, 1965,initscurrentversion,andpermissionforusemustalwaysbeobtainedfromSpringer.Violations areliabletoprosecutionundertheGermanCopyrightLaw. Theuseofgeneraldescriptivenames,registerednames,trademarks,etc.inthispublicationdoesnot imply, even in the absence of a specific statement, that such names are exempt from the relevant protectivelawsandregulationsandthereforefreeforgeneraluse. Coverdesign:SPiPublisherServices Printedonacid-freepaper SpringerispartofSpringerScience+BusinessMedia(www.springer.com) Preface Methanogens are prokaryotic microorganisms that produce methane as an end- product of their metabolism. They are strictly anaerobic archaea belonging to the taxon Euryarchaeota. Methanogens occupy a wide variety of anaerobic environ- ments, even extreme habitats characterized by high temperature, salinity, and extreme pH (Liu and Whitman 2008). More temperate habitats include marine and freshwater sediments, flooded soils, landfills, anaerobic digesters, geothermal systems, and the heartwood of trees. Notably, methanogens also thrive in the cytoplasm of anaerobic unicellular eukaryotes and in the gastrointestinal tracts of animalsandhumans.Frequently,theyattachtotheinternalsurfacesofthegastro- intestinal tracts and of the protists living in intestinal environments with the aid ofspecialadhesion-likeproteins(LiuandWhitman2008).Ininsectguts,methano- gens have to cope with a very special environment, since they are exposed to a continuousinfluxofoxygenthroughthegutwallthatchallengesthestrictlyanaero- bic symbionts. Notwithstanding that methanogens are very diverse, they can use only a very restricted number of substrates. They are unable to use organic substances (with the exception of acetate and formate), and consequently, methanogens must rely on CO and methyl-group containing compounds and 2 acetate, which are provided by the fermentations performed by complex anaerobic bacterial communities. The methanogenic substrates are predominantly metabo- lized with the aid of H that is provided by syntrophic bacterial communities or, 2 in the case of certain anaerobic protists, by the action of specialized mitochondri- on-derived organelles, the hydrogenosomes. Since both hydrogenosomes and syntrophic bacterial communities rely on a low concentration of H , the interspe- 2 cies hydrogen transfer that is mediated by methanogens is crucial for the proper functioning of hydrogenosomes and syntrophic anaerobic bacterial communities. The synthesis of methane follows a complex biochemical pathway that is characterized by a number of unique coenzymes and membrane-bound enzyme complexes.IthasbeenreviewedrecentlybyHedderichandWhitman(2006).CO 2 isreducedtomethanebyH inhydrogenotrophicmethanogens,whichrepresentthe 2 majority of the methanogens living in symbiosis with protists and multicellular animals. The second type of substrate, methylgroup containing compounds v vi Preface includingmethanol,methylatedamines,andmethylatedsulfides,isusedbymethy- lotrophicmethanogens,whicharepredominantlyfoundinthegastrointestinaltracts ofmammalsandinsects.Thethirdtypeofsubstrateisacetate,whichismetabolized by acetoclastic methanogens. Although only two genera (Methanosarcina and Methanosaeta) use acetate as substrate, as much as two-thirds of the biologically generatedmethaneisderivedfromacetate.Notably,acetoclasticmethanogensare rare among the symbiotic methanogens. This is not surprising since symbiotic acetoclastic methanogens compete with their hosts for acetate. Notwithstanding, an acetoclastic methanogen has recently been identified as endosymbiont in the free-livingprotistMetopuses(Narayananetal.2009). Methanogens are abundant in habitats where electron acceptors such as O , 2 NO (cid:1), Fe3+, and SO 2(cid:1) are limiting (Liu and Whitman 2008). In such methano- 3 4 genic habitats, complex organic matter is degraded to methane by the syntrophic action of different groups of anaerobic bacteria. Organic polymers are degraded initially by specialized bacteria to sugars, lactate, short-chain fatty acids, and alcohols.Theseproductsarefurtherfermentedbyotherbacteriatoacetate,formate, H and CO , which are the substrates used by methanogens. These methanogens 2, 2 catalyze the terminal step in the anaerobic food chain by converting the various methanogenicsubstrates tomethane,whichisreleasedintothe atmosphere.More than70%oftheannualglobalmethaneemission(rangingfrom500to600TgCH / 4 year) stem from biological methanogenesis and contribute significantly to global warming.(Whitmanetal.2006).Thesymbioticmethanogensinthegastrointestinal tract of ruminants and other “methanogenic” mammals contribute significantly to the global methane budget. Especially the rumen hosts an impressive diversity of methanogens,whichhavebeenstudiedusingculture-independent16SrRNAmeth- ods(JanssenandKirs2008;Wrightetal.2004).Insects,particularlytermites,also hostverycomplexmethanogeniccommunities,buttheyreleasemuchlessmethane duetotheconcomitantoxidationofmethaneinthesoilandthetermitemounds.On the other hand, the contributions by the methanogenic endosymbionts of protists livinginfreshwatersedimentscanbeneglected(vanHoeketal.2006). This monograph deals with methanogenic endosymbionts of free-living and symbiotic protists, episymbionts of rumen ciliates, methanogenic endosymbionts of ciliates, and termite flagellates, which are accompanied by bacterial endosym- bionts,andmethanogensinthegastrointestinaltractofvertebratesandarthropods. Onereviewsummarizesourknowledgeaboutthegenomicconsequencesofliving together in symbiotic associations; another review discusses the role of methano- gens in syntrophic degradation. Finally, the current state of information about hydrogenosomeshasbeenreviewed. Wegratefullyacknowledgetheeffortsoftheauthorswhocontributedexcellent chapters to this volume of Microbiology Monographs. We thank Springer for publishingthismonographandJuttaLindenbornforsupport. Nijmegen,TheNetherlands JohannesH.P.Hackstein Mu¨nster,Germany AlexanderSteinbu¨chel Preface vii References HedderichR,WhitmanWB(2006)Physiologyandbiochemistryofmethane-producingArchaea. In:DworkinMetal(eds)Theprokaryotes,vol2,3rdedn.Springer,NewYork,pp1050–1079 Janssen PH, Kirs M (2008) Structure of the archaeal community of the rumen. Appl Environ Microbiol74:3619–3625 LiuY,WhitmanWB(2008)Metabolic,phylogenetic,andecologicaldiversityofthemethano- genicarchaea.AnnNYAcadSci1125:171–189 NarayananN,KrishnakumarB,AnupamaVN,ManilalVB(2009)Methanosaetasp.,themajor archaealendosymbiontofMetopuses.ResMicrobiol160:600–607 vanHoekAHAM,vanAlenTA,VogelsGD,HacksteinJHP(2006)Contributionbythemethano- genic endosymbionts of anaerobic ciliates to methane production in Dutch freshwater sedi- ments.ActaProtozool45:215–224 WhitmanWB,BowenT,BooneD(2006)Themethanogenicbacteria.In:DworkinMetal(eds) Theprokaryotes,vol.3,3rdedn.Springer,NewYork,pp165–207 Wright ADG, Williams AJ, Winder B, Christophersen CT, Rodgers SL, Smith KD (2004) Molecular diversity ofrumen methanogens from sheep in Western Australia. Appl Environ Microbiol70:1263–1270 . Contents Free-LivingProtozoawithEndosymbioticMethanogens ................... 1 TomFenchelandBlandJ.Finlay AnaerobicCiliatesandTheirMethanogenicEndosymbionts ............. 13 JohannesH.P.Hackstein SymbioticMethanogensandRumenCiliates ............................... 25 KazunariUshida TheMethanogenicandEubacterialEndosymbiontsofTrimyema ........ 35 NaoyaShinzatoandYoichiKamagata TermiteGutFlagellatesandTheirMethanogenic andEubacterialSymbionts ................................................... 55 YuichiHongohandMoriyaOhkuma MethanogensintheDigestiveTractofTermites ........................... 81 AndreasBrune MethanogenicArchaeainHumansandOtherVertebrates .............. 101 EverlyConwaydeMacarioandAlbertoJ.L.Macario MethanogensintheGastro-IntestinalTractofAnimals .................. 115 JohannesH.P.HacksteinandTheoA.vanAlen SyntrophyinMethanogenicDegradation .................................. 143 PetraWorm,NicolaiMu¨ller,CarolineM.Plugge,AlfonsJ.M.Stams, andBernhardSchink ix
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