Microorganism

25/03/2015
MicroorganismWikipedia,thefreeencyclopedia
Microorganism
FromWikipedia,thefreeencyclopedia
Amicroorganism(fromtheGreek:,mikros,"small"
and,organisms,"organism")isamicroscopic
livingorganism,whichmaybesinglecelled[1]ormulticellular.
Thestudyofmicroorganismsiscalledmicrobiology,asubject
thatbeganwiththediscoveryofmicroorganismsin1674by
AntonievanLeeuwenhoek,usingamicroscopeofhisown
design.
Microorganismsareverydiverseandincludeallthebacteria
andarchaeaandalmostalltheprotozoa.Theyalsoinclude
somefungi,algae,andcertainanimals,suchasrotifers.Many
macroscopicanimalsandplantshavemicroscopicjuvenile
stages.Somemicrobiologistsalsoclassifyviruses(andviroids)
asmicroorganisms,butothersconsidertheseasnonliving.[2][3]
AclusterofEscherichiacolibacteria
magnified10,000times
Microorganismsliveineverypartofthebiosphere,includingsoil,hotsprings,"sevenmilesdeep"inthe
ocean,"40mileshigh"intheatmosphereandinsiderocksfardownwithintheEarth'scrust(seealso
endolith).[4]Microorganisms,undercertaintestconditions,havebeenobservedtothriveinthevacuumof
outerspace.[5][6]Thetotalamountofsoilandsubsurfacebacterialcarbonisestimatedas5x1017g,orthe
"weightoftheUnitedKingdom".[4]Themassofprokaryotemicroorganismswhichincludesbacteriaand
archaea,butnotthenucleatedeukaryotemicroorganismsmaybeasmuchas0.8trilliontonsofcarbon
(ofthetotalbiospheremassof4trilliontons).[7]On17March2013,researchersreporteddatathat
suggestedmicrobiallifeformsthriveintheMarianaTrench.thedeepestspotintheEarth'soceans.[8][9]
Otherresearchersreportedrelatedstudiesthatmicroorganismsthriveinsiderocksupto580m(1,900ft
0.36mi)belowtheseafloorunder2,590m(8,500ft1.61mi)ofoceanoffthecoastofthenorthwestern
UnitedStates[8][10]aswellas2,400m(7,900ft1.5mi)beneaththeseabedoffJapan.[11]On20August
2014,scientistsconfirmedtheexistenceofmicroorganismsliving800m(2,600ft0.50mi)belowtheice
ofAntarctica.[12][13]Accordingtooneresearcher,"Youcanfindmicrobeseverywherethey'reextremely
adaptabletoconditions,andsurvivewherevertheyare."[8]
Microorganismsarecrucialtonutrientrecyclinginecosystemsastheyactasdecomposers.Assome
microorganismscanfixnitrogen,theyareavitalpartofthenitrogencycle,andrecentstudiesindicatethat
airbornemicroorganismsmayplayaroleinprecipitationandweather.[14]Microorganismsarealso
exploitedinbiotechnology,bothintraditionalfoodandbeveragepreparation,andinmoderntechnologies
basedongeneticengineering.Asmallproportionofmicroorganismsarepathogenicandcausediseaseand
evendeathinplantsandanimals.[15]Microorganismsareoftenreferredtoasmicrobes,butthisisusually
usedinreferencetopathogens.
Contents
http://en.wikipedia.org/wiki/Microorganism
1/21
25/03/2015
1Evolution
2Premicrobiology
3Historyofmicroorganisms'discovery
4Classificationandstructure
4.1Prokaryotes
4.1.1Bacteria
4.1.2Archaea
4.2Eukaryotes
4.2.1Protists
4.2.2Animals
4.2.3Fungi
4.2.4Plants
5Habitatsandecology
5.1Extremophiles
5.2Soilmicroorganisms
5.3Symbioticmicroorganisms
6Importance
6.1Useindigestion
6.2Useinfoodproduction
6.3Useinwatertreatment
6.4Useinenergy
6.5Useinproductionofchemicals,enzymesetc.
6.6Useinscience
6.7Useinwarfare
7Importanceinhumanhealth
7.1Humandigestion
7.2Diseasescausedbymicrobes
8Importanceinecology
9Hygiene
10Seealso
11References
12Externallinks
Evolution
2/21
25/03/2015
SinglecelledmicroorganismswerethefirstformsoflifetodeveloponEarth,approximately34billion
yearsago.[16][17][18]Furtherevolutionwasslow,[19]andforabout3billionyearsinthePrecambrianeon,all
organismsweremicroscopic.[20]So,formostofthehistoryoflifeonEarth,theonlyformsoflifewere
microorganisms.[21]Bacteria,algaeandfungihavebeenidentifiedinamberthatis220millionyearsold,
whichshowsthatthemorphologyofmicroorganismshaschangedlittlesincetheTriassicperiod.[22]The
newlydiscoveredbiologicalroleplayedbynickel,howeverespeciallythatengenderedbyvolcanic
eruptionsfromtheSiberianTraps(siteofthemoderncityofNorilsk)isthoughttohaveacceleratedthe
evolutionofmethanogenstowardstheendofthePermianTriassicextinctionevent.[23]
Microorganismstendtohavearelativelyfastrateofevolution.Mostmicroorganismscanreproduce
rapidly,andbacteriaarealsoabletofreelyexchangegenesthroughconjugation,transformationand
transduction,evenbetweenwidelydivergentspecies.[24]Thishorizontalgenetransfer,coupledwithahigh
mutationrateandmanyothermeansofgeneticvariation,allowsmicroorganismstoswiftlyevolve(via
naturalselection)tosurviveinnewenvironmentsandrespondtoenvironmentalstresses.Thisrapid
evolutionisimportantinmedicine,asithasledtotherecentdevelopmentof"superbugs",pathogenic
bacteriathatareresistanttomodernantibiotics.[25]
Premicrobiology
Thepossibilitythatmicroorganismsexistwasdiscussedformanycenturiesbeforetheirdiscoveryinthe
17thcentury.TheexistenceofunseenmicrobiologicallifewaspostulatedbyJainism,whichisbasedon
Mahavira'steachingsasearlyas6thcenturyBCE.[26]PaulDundasnotesthatMahaviraassertedthe
existenceofunseenmicrobiologicalcreatureslivinginearth,water,airandfire.[27]TheJainscripturesalso
describenigodas,whicharesubmicroscopiccreatureslivinginlargeclustersandhavingaveryshortlife,
whicharesaidtopervadeeverypartoftheuniverse,eventhetissuesofplantsandanimals.[28]Theearliest
knownideatoindicatethepossibilityofdiseasesspreadingbyyetunseenorganismswasthatoftheRoman
scholarMarcusTerentiusVarroina1stcenturyBCbooktitledOnAgricultureinwhichhewarnsagainst
locatingahomesteadnearswamps:
andbecausetherearebredcertainminutecreaturesthatcannotbeseenbytheeyes,which
floatintheairandenterthebodythroughthemouthandnoseandtheycauseserious
diseases.[29]
InTheCanonofMedicine(1020),AbAlibnSn(Avicenna)hypothesizedthattuberculosisandother
diseasesmightbecontagious[30][31]
In1546,GirolamoFracastoroproposedthatepidemicdiseaseswerecausedbytransferableseedlikeentities
thatcouldtransmitinfectionbydirectorindirectcontact,orevenwithoutcontactoverlongdistances.
Alltheseearlyclaimsabouttheexistenceofmicroorganismswerespeculativeandwhilegroundedon
indirectobservations,theywerenotbasedondirectobservationofmicroorganismsorsystematized
empiricalinvestigation,e.g.experimentation.Microorganismswereneitherproven,observed,nor
accuratelydescribeduntilthe17thcentury.Thereasonforthiswasthatalltheseearlystudieslackedthe
microscope.
3/21
25/03/2015
Historyofmicroorganisms'discovery
AntonieVanLeeuwenhoek(16321723)
wasoneofthefirstpeopletoobserve
microorganisms,usingmicroscopesofhis
owndesign.[32]RobertHooke,a
contemporaryofLeeuwenhoek,alsoused
microscopestoobservemicrobiallifehis
1665bookMicrographiadescribesthese
observationsandcoinedthetermcell.
AntonievanLeeuwenhoek,
thefirstmicrobiologistand
thefirsttoobserve
microorganismsusinga
microscope
LouisPasteurshowedthat
Spallanzani'sfindingsheld
evenifaircouldenter
throughafilterthatkept
particlesout
BeforeLeeuwenhoek'sdiscoveryof
microorganismsin1675,ithadbeena
mysterywhygrapescouldbeturnedinto
wine,milkintocheese,orwhyfoodwould
spoil.Leeuwenhoekdidnotmakethe
connectionbetweentheseprocessesand
microorganisms,butusingamicroscope,he
didestablishthattherewereformsoflife
thatwerenotvisibletothenaked
eye.[33][34]Leeuwenhoek's
discovery,alongwithsubsequent
observationsbySpallanzaniand
Pasteur,endedthelongheldbelief
thatlifespontaneouslyappeared
fromnonlivingsubstancesduring
theprocessofspoilage.
LazzaroSpallanzani(17291799)
foundthatboilingbrothwould
steriliseit,killingany
microorganismsinit.Healsofound
thatnewmicroorganismscouldonly
settleinabrothifthebrothwas
exposedtoair.
LazzaroSpallanzanishowed
thatboilingabrothstopped
itfromdecaying
RobertKochshowedthat
microorganismscauseddisease
LouisPasteur(18221895)
expandeduponSpallanzani'sfindingsbyexposingboiledbrothsto
theair,invesselsthatcontainedafiltertopreventallparticlesfrompassingthroughtothegrowthmedium,
andalsoinvesselswithnofilteratall,withairbeingadmittedviaacurvedtubethatwouldnotallowdust
particlestocomeincontactwiththebroth.Byboilingthebrothbeforehand,Pasteurensuredthatno
microorganismssurvivedwithinthebrothsatthebeginningofhisexperiment.Nothinggrewinthebroths
inthecourseofPasteur'sexperiment.Thismeantthatthelivingorganismsthatgrewinsuchbrothscame
fromoutside,assporesondust,ratherthanspontaneouslygeneratedwithinthebroth.Thus,Pasteurdealt
thedeathblowtothetheoryofspontaneousgenerationandsupportedgermtheory.
4/21
25/03/2015
In1876,RobertKoch(18431910)establishedthatmicroorganismscancausedisease.Hefoundthatthe
bloodofcattlewhichwereinfectedwithanthraxalwayshadlargenumbersofBacillusanthracis.Koch
foundthathecouldtransmitanthraxfromoneanimaltoanotherbytakingasmallsampleofbloodfromthe
infectedanimalandinjectingitintoahealthyone,andthiscausedthehealthyanimaltobecomesick.He
alsofoundthathecouldgrowthebacteriainanutrientbroth,theninjectitintoahealthyanimal,andcause
illness.Basedontheseexperiments,hedevisedcriteriaforestablishingacausallinkbetweena
microorganismandadiseaseandthesearenowknownasKoch'spostulates.[35]Althoughthesepostulates
cannotbeappliedinallcases,theydoretainhistoricalimportancetothedevelopmentofscientificthought
andarestillbeingusedtoday.[36]
On8November2013,scientistsreportedthediscoveryofwhatmaybetheearliestsignsoflifeonEarth
theoldestcompletefossilsofamicrobialmat(associatedwithsandstoneinWesternAustralia)estimatedto
be3.48billionyearsold.[37][38]
Classificationandstructure
Microorganismscanbefoundalmost
anywhereinthetaxonomicorganizationof
lifeontheplanet.Bacteriaandarchaeaare
almostalwaysmicroscopic,whilea
numberofeukaryotesarealsomicroscopic,
includingmostprotists,somefungi,aswell
assomeanimalsandplants.Virusesare
generallyregardedasnotlivingand
thereforenotconsideredas
microorganisms,althoughthefieldof
microbiologyalsoencompassesthestudy
ofviruses.
Prokaryotes
Prokaryotesareorganismsthatlackacell
nucleusandtheothermembranebound
organelles.Theyarealmostalways
unicellular,althoughsomespeciessuchas
myxobacteriacanaggregateintocomplex
structuresaspartoftheirlifecycle.
Evolutionarytreeshowingthecommonancestryofallthree
domainsoflife. [39]Bacteriaarecoloredblue,eukaryotesred,and
archaeagreen.Relativepositionsofsomephylaareshownaround
thetree.
Consistingoftwodomains,bacteriaandarchaea,theprokaryotesarethemostdiverseandabundantgroup
oforganismsonEarthandinhabitpracticallyallenvironmentswherethetemperatureisbelow+140C.
Theyarefoundinwater,soil,air,animals'gastrointestinaltracts,hotspringsandevendeepbeneaththe
Earth'scrustinrocks.[40]Practicallyallsurfacesthathavenotbeenspeciallysterilizedarecoveredby
prokaryotes.ThenumberofprokaryotesonEarthisestimatedtobearoundfivemilliontrilliontrillion,or5
1030,accountingforatleasthalfthebiomassonEarth.[41]
Bacteria
5/21
25/03/2015
Almostallbacteriaareinvisibletothenakedeye,withafew
extremelyrareexceptions,suchasThiomargarita
namibiensis.[42]Theylackanucleusandothermembrane
boundorganelles,andcanfunctionandreproduceasindividual
cells,butoftenaggregateinmulticellularcolonies.[43]Their
genomeisusuallyasingleloopofDNA,althoughtheycanalso
harborsmallpiecesofDNAcalledplasmids.Theseplasmids
canbetransferredbetweencellsthroughbacterialconjugation.
Bacteriaaresurroundedbyacellwall,whichprovidesstrength
andrigiditytotheircells.Theyreproducebybinaryfissionor
sometimesbybudding,butdonotundergomeioticsexual
Staphylococcusaureusbacteriamagnified
reproduction.However,manybacterialspeciescantransfer
about10,000x
DNAbetweenindividualcellsbyaprocessreferredtoas
[44][45]
naturaltransformation.
Innature,thedevelopmentof
competencefortransformationisusuallyassociatedwithstressfulenvironmentalconditions,andseemsto
beanadaptationforfacilitatingrepairofDNAdamageinrecipientcells.[46][47]Somespeciesform
extraordinarilyresilientspores,butforbacteriathisisamechanismforsurvival,notreproduction.Under
optimalconditionsbacteriacangrowextremelyrapidlyandcandoubleasquicklyasevery20minutes.[48]
Archaea
Archaeaarealsosinglecelledorganismsthatlacknuclei.Inthepast,thedifferencesbetweenbacteriaand
archaeawerenotrecognisedandarchaeawereclassifiedwithbacteriaaspartofthekingdomMonera.
However,in1990themicrobiologistCarlWoeseproposedthethreedomainsystemthatdividedliving
thingsintobacteria,archaeaandeukaryotes.[49]Archaeadifferfrombacteriainboththeirgeneticsand
biochemistry.Forexample,whilebacterialcellmembranesaremadefromphosphoglycerideswithester
bonds,archaeanmembranesaremadeofetherlipids.[50]
Archaeawereoriginallydescribedinextremeenvironments,suchashotsprings,buthavesincebeenfound
inalltypesofhabitats.[51]Onlynowarescientistsbeginningtorealizehowcommonarchaeaareinthe
environment,withcrenarchaeotabeingthemostcommonformoflifeintheocean,dominatingecosystems
below150mindepth.[52][53]Theseorganismsarealsocommoninsoilandplayavitalroleinammonia
oxidation.[54]
Eukaryotes
Mostlivingthingsthatarevisibletothenakedeyeintheiradultformareeukaryotes,includinghumans.
However,alargenumberofeukaryotesarealsomicroorganisms.Unlikebacteriaandarchaea,eukaryotes
containorganellessuchasthecellnucleus,theGolgiapparatusandmitochondriaintheircells.Thenucleus
isanorganellethathousestheDNAthatmakesupacell'sgenome.DNAitselfisarrangedincomplex
chromosomes.[55]Mitochondriaareorganellesvitalinmetabolismastheyarethesiteofthecitricacid
cycleandoxidativephosphorylation.Theyevolvedfromsymbioticbacteriaandretainaremnant
genome.[56]Likebacteria,plantcellshavecellwalls,andcontainorganellessuchaschloroplastsinaddition
totheorganellesinothereukaryotes.Chloroplastsproduceenergyfromlightbyphotosynthesis,andwere
alsooriginallysymbioticbacteria.[56]
6/21
25/03/2015
Unicellulareukaryotesconsistofasinglecellthroughouttheirlifecycle.Thisqualificationissignificant
sincemostmulticellulareukaryotesconsistofasinglecellcalledazygoteonlyatthebeginningoftheirlife
cycles.Microbialeukaryotescanbeeitherhaploidordiploid,andsomeorganismshavemultiplecell
nuclei.[57]
Unicellulareukaryotesusuallyreproduceasexuallybymitosisunderfavorableconditions.However,under
stressfulconditionssuchasnutrientlimitationsandotherconditionsassociatedwithDNAdamage,they
tendtoreproducesexuallybymeiosisandsyngamy.[46][58]
Protists
Ofeukaryoticgroups,theprotistsaremostcommonlyunicellularandmicroscopic.Thisisahighlydiverse
groupoforganismsthatarenoteasytoclassify.[59][60]Severalalgaespeciesaremulticellularprotists,and
slimemoldshaveuniquelifecyclesthatinvolveswitchingbetweenunicellular,colonial,andmulticellular
forms.[61]Thenumberofspeciesofprotistsisunknownsincewemayhaveidentifiedonlyasmallportion.
Studiesfrom20012004haveshownthatahighdegreeofprotistdiversityexistsinoceans,deepseavents,
riversedimentandanacidicriverwhichsuggeststhatalargenumberofeukaryoticmicrobialcommunities
haveyettobediscovered.[62][63]
Animals
Somemicroanimalsaremulticellularbutatleastoneanimalgroup,
Myxozoa,isunicellularinitsadultform.Microscopicarthropods
includedustmitesandspidermites.Microscopiccrustaceansinclude
copepods,somecladoceraandwaterbears.Manynematodesarealso
toosmalltobeseenwiththenakedeye.Acommongroupof
microscopicanimalsaretherotifers,whicharefilterfeedersthatare
usuallyfoundinfreshwater.Somemicroanimalsreproduceboth
sexuallyandasexuallyandmayreachnewhabitatsbyproducingeggs
whichcansurviveharshenvironmentsthatwouldkilltheadult
animal.However,somesimpleanimals,suchasrotifers,tardigrades
andnematodes,candryoutcompletelyandremaindormantforlong
periodsoftime.[64]
AmicroscopicmiteLorryiaformosa
Fungi
Thefungihaveseveralunicellularspecies,suchasbaker'syeast(Saccharomycescerevisiae)andfission
yeast(Schizosaccharomycespombe).Somefungi,suchasthepathogenicyeastCandidaalbicans,can
undergophenotypicswitchingandgrowassinglecellsinsomeenvironments,andfilamentoushyphaein
others.[65]Fungireproducebothasexually,bybuddingorbinaryfission,aswellbyproducingspores,which
arecalledconidiawhenproducedasexually,orbasidiosporeswhenproducedsexually.
Plants
7/21
25/03/2015
Thegreenalgaearealargegroupofphotosyntheticeukaryotesthatincludemanymicroscopicorganisms.
Althoughsomegreenalgaeareclassifiedasprotists,otherssuchascharophytaareclassifiedwith
embryophyteplants,whicharethemostfamiliargroupoflandplants.Algaecangrowassinglecells,orin
longchainsofcells.Thegreenalgaeincludeunicellularandcolonialflagellates,usuallybutnotalways
withtwoflagellapercell,aswellasvariouscolonial,coccoid,andfilamentousforms.IntheCharales,
whicharethealgaemostcloselyrelatedtohigherplants,cellsdifferentiateintoseveraldistincttissues
withintheorganism.Thereareabout6000speciesofgreenalgae.[66]
Habitatsandecology
Microorganismsarefoundinalmosteveryhabitatpresentinnature.Eveninhostileenvironmentssuchas
thepoles,deserts,geysers,rocks,andthedeepsea.Sometypesofmicroorganismshaveadaptedtothe
extremeconditionsandsustainedcoloniestheseorganismsareknownasextremophiles.Extremophiles
havebeenisolatedfromrocksasmuchas7kilometresbelowtheEarth'ssurface,[67]andithasbeen
suggestedthattheamountoflivingorganismsbelowtheEarth'ssurfacemaybecomparablewiththe
amountoflifeonorabovethesurface.[40]Extremophileshavebeenknowntosurviveforaprolongedtime
inavacuum,andcanbehighlyresistanttoradiation,whichmayevenallowthemtosurviveinspace.[68]
Manytypesofmicroorganismshaveintimatesymbioticrelationshipswithotherlargerorganismssomeof
whicharemutuallybeneficial(mutualism),whileotherscanbedamagingtothehostorganism(parasitism).
Ifmicroorganismscancausediseaseinahosttheyareknownaspathogensandthentheyaresometimes
referredtoasmicrobes.
Extremophiles
Extremophilesaremicroorganismsthathaveadaptedsothattheycansurviveandeventhriveinconditions
thatarenormallyfataltomostlifeforms.Forexample,somespecieshavebeenfoundinthefollowing
extremeenvironments:
Temperature:ashighas130C(266F),[69]aslowas17C(1F)[70]
Acidity/alkalinity:lessthanpH0,[71]uptopH11.5[72]
Salinity:uptosaturation[73]
Pressure:upto1,0002,000atm,downto0atm(e.g.vacuumofspace)[74]
Radiation:upto5kGy[75]
Extremophilesaresignificantindifferentways.TheyextendterrestriallifeintomuchoftheEarth's
hydrosphere,crustandatmosphere,theirspecificevolutionaryadaptationmechanismstotheirextreme
environmentcanbeexploitedinbiotechnology,andtheirveryexistenceundersuchextremeconditions
increasesthepotentialforextraterrestriallife.[76]
Soilmicroorganisms
Thenitrogencycleinsoilsdependsonthefixationofatmosphericnitrogen.Onewaythiscanoccurisin
thenodulesintherootsoflegumesthatcontainsymbioticbacteriaofthegeneraRhizobium,
Mesorhizobium,Sinorhizobium,Bradyrhizobium,andAzorhizobium.[77]
8/21
25/03/2015
Symbioticmicroorganisms
Symbioticmicroorganismssuchasfungiandalgaeformanassociationinlichen.Certainfungiform
mycorrhizalsymbioseswithtreesthatincreasethesupplyofnutrientstothetree.
Importance
Microorganismsarevitaltohumansandtheenvironment,astheyparticipateinthecarbonandnitrogen
cycles,aswellasfulfillingothervitalrolesinvirtuallyallecosystems,suchasrecyclingotherorganisms'
deadremainsandwasteproductsthroughdecomposition.Microorganismsalsohaveanimportantplacein
mosthigherordermulticellularorganismsassymbionts.ManyblamethefailureofBiosphere2onan
improperbalanceofmicroorganisms.[78]
Useindigestion
Someformsofbacteriathatliveinanimals'stomachshelpintheirdigestion.Forexample,cowshavea
varietyofdifferentmicroorganismsintheirstomachsthatareessentialintheirdigestionofgrassandhay.
Thegastrointestinaltractcontainsanimmenselycomplexecologyofmicroorganisms.Atypicalperson
harborsmorethan500distinctspeciesofbacteria,representingdozensofdifferentlifestylesand
capabilities.Thecompositionanddistributionofthismenagerievarieswithage,stateofhealthanddiet.
Thenumberandtypeofbacteriainthegastrointestinaltractvarydramaticallybyregion.Inhealthy
individualsthestomachandproximalsmallintestinecontainfewmicroorganisms,largelyaresultofthe
bacteriocidalactivityofgastricacidthosethatarepresentareaerobesandfacultativeanaerobes.One
interestingtestimonytotheabilityofgastricacidtosuppressbacterialpopulationsisseeninpatientswith
achlorhydria,ageneticconditionwhichpreventssecretionofgastricacid.Suchpatients,whichare
otherwisehealthy,mayhaveasmanyas10,000to100,000,000microorganismspermlofstomach
contents.
Insharpcontrasttothestomachandsmallintestine,thecontentsofthecolonliterallyteemwithbacteria,
predominantlystrictanaerobes(bacteriathatsurviveonlyinenvironmentsvirtuallydevoidofoxygen).
Betweenthesetwoextremesisatransitionalzone,usuallyintheileum,wheremoderatenumbersofboth
aerobicandanaerobicbacteriaarefound.
Thegastrointestinaltractissterileatbirth,butcolonizationtypicallybeginswithinafewhoursofbirth,
startinginthesmallintestineandprogressingcaudallyoveraperiodofseveraldays.Inmostcircumstances,
a"mature"microbialfloraisestablishedby3to4weeksofage.
Itisalsoclearthatmicrobialpopulationsexertaprofoundeffectonstructureandfunctionofthedigestive
tract.Forexample:
Themorphologyoftheintestineofgermfreeanimalsdiffersconsiderablyfromnormalanimalsvilliof
thesmallintestineareremarkablyregular,therateofepithelialcellrenewisreducedand,asonewould
expect,thenumberandsizeofPeyer'spatchesisreduced.Thececumofgermfreeratsisroughly10times
thesizeofthatinaconventionalrat.Bacteriaintheintestinallumenmetabolizeavarietyofsterolsand
steroids.Forexample,bacteriaconvertthebilesaltcholicacidtodeoxycholicacid.Smallintestinalbacteria
alsohaveanimportantroleinsexsteroidmetabolism.Finally,bacterialpopulationsinthelargeintestine
9/21
25/03/2015
digestcarbohydrates,proteinsandlipidsthatescapedigestionandabsorptioninsmallintestine.This
fermentation,particularlyofcellulose,isofcriticalimportancetoherbivoreslikecattleandhorseswhich
makealivingbyconsumingplants.However,itseemsthatevenspecieslikehumansandrodentsderive
significantbenefitfromthenutrientsliberatedbyintestinalmicroorganisms.
Useinfoodproduction
Microorganismsareusedinbrewing,winemaking,baking,picklingandotherfoodmakingprocesses.
Theyarealsousedtocontrolthefermentationprocessintheproductionofcultureddairyproductssuchas
yogurtandcheese.Theculturesalsoprovideflavourandaroma,andinhibitundesirableorganisms.[79]
Useinwatertreatment
Themajorityofalloxidativesewagetreatmentprocessesrelyonalargerangeofmicroorganismstooxidise
organicconstituentswhicharenotamenabletosedimentationorflotation.Anaerobicmicroorganismsare
alsousedtoreducesludgesolidsproducingmethanegas(amongstothergases)andasterilemineralised
residue.Inpotablewatertreatment,onemethod,theslowsandfilter,employsacomplexgelatinouslayer
composedofawiderangeofmicroorganismstoremovebothdissolvedandparticulatematerialfromraw
water.[80]
Useinenergy
Microorganismsareusedinfermentationtoproduceethanol,[81]andinbiogasreactorstoproduce
methane.[82]Scientistsareresearchingtheuseofalgaetoproduceliquidfuels,[83]andbacteriatoconvert
variousformsofagriculturalandurbanwasteintousablefuels.[84]
Useinproductionofchemicals,enzymesetc.
Microorganismsareusedformanycommercialandindustrialproductionofchemicals,enzymesandother
bioactivemolecules.
Examplesoforganicacidproducedinclude
Aceticacid:ProducedbythebacteriumAcetobacteracetiandotheraceticacidbacteria(AAB)
Butyricacid(butanoicacid):ProducedbythebacteriumClostridiumbutyricum
Lacticacid:Lactobacillusandotherscommonlycalledaslacticacidbacteria(LAB)
Citricacid:ProducedbythefungusAspergillusniger
Microorganismsareusedforpreparationofbioactivemoleculesandenzymes.
StreptokinaseproducedbythebacteriumStreptococcusandmodifiedbygeneticengineeringisused
asaclotbusterforremovingclotsfromthebloodvesselsofpatientswhohaveundergonemyocardial
infarctionsleadingtoheartattack.
CyclosporinAisabioactivemoleculeusedasanimmunosuppressiveagentinorgantransplantation
10/21
25/03/2015
StatinsproducedbytheyeastMonascuspurpureusarecommercialisedasbloodcholesterollowering
agentswhichactbycompetitivelyinhibitingtheenzymeresponsibleforsynthesisofcholesterol.[85]
Useinscience
Microorganismsareessentialtoolsinbiotechnology,biochemistry,genetics,andmolecularbiology.The
yeasts(Saccharomycescerevisiae)andfissionyeast(Schizosaccharomycespombe)areimportantmodel
organismsinscience,sincetheyaresimpleeukaryotesthatcanbegrownrapidlyinlargenumbersandare
easilymanipulated.[86]Theyareparticularlyvaluableingenetics,genomicsandproteomics.[87][88]
Microorganismscanbeharnessedforusessuchascreatingsteroidsandtreatingskindiseases.Scientistsare
alsoconsideringusingmicroorganismsforlivingfuelcells,[89]andasasolutionforpollution.[90]
Useinwarfare
IntheMiddleAges,diseasedcorpseswerethrownintocastlesduringsiegesusingcatapultsorothersiege
engines.Individualsnearthecorpseswereexposedtothepathogenandwerelikelytospreadthatpathogen
toothers.[91]
Importanceinhumanhealth
Humandigestion
Microorganismscanformanendosymbioticrelationshipwithother,largerorganisms.Forexample,the
bacteriathatlivewithinthehumandigestivesystemcontributetogutimmunity,synthesisevitaminssuchas
folicacidandbiotin,andfermentcomplexindigestiblecarbohydrates.[92]
Diseasescausedbymicrobes
Microorganismsarethecauseofmanyinfectiousdiseases.Theorganismsinvolvedincludepathogenic
bacteria,causingdiseasessuchasplague,tuberculosisandanthraxprotozoa,causingdiseasessuchas
malaria,sleepingsickness,dysenteryandtoxoplasmosisandalsofungicausingdiseasessuchasringworm,
candidiasisorhistoplasmosis.However,otherdiseasessuchasinfluenza,yellowfeverorAIDSarecaused
bypathogenicviruses,whicharenotusuallyclassifiedaslivingorganismsandarenot,therefore,
microorganismsbythestrictdefinition.Asof2007,noclearexamplesofarchaeanpathogensare
known,[93]althougharelationshiphasbeenproposedbetweenthepresenceofsomearchaeanmethanogens
andhumanperiodontaldisease.[94]
Importanceinecology
MicroorganismsplaycriticalrolesinEarth'sbiogeochemicalcyclesastheyareresponsiblefor
decompositionandplayvitalpartsincarbonandnitrogenfixation,aswellasoxygenproduction.
Hygiene
11/21
25/03/2015
Hygieneistheavoidanceofinfectionorfoodspoilingbyeliminatingmicroorganismsfromthe
surroundings.Asmicroorganisms,inparticularbacteria,arefoundvirtuallyeverywhere,thelevelsof
harmfulmicroorganismscanbereducedtoacceptablelevels.However,insomecases,itisrequiredthatan
objectorsubstancebecompletelysterile,i.e.devoidofalllivingentitiesandviruses.Agoodexampleof
thisisahypodermicneedle.
Infoodpreparationmicroorganismsarereducedbypreservationmethods(suchastheadditionofvinegar),
cleanutensilsusedinpreparation,shortstorageperiods,orbycooltemperatures.Ifcompletesterilityis
needed,thetwomostcommonmethodsareirradiationandtheuseofanautoclave,whichresemblesa
pressurecooker.
Thereareseveralmethodsforinvestigatingthelevelofhygieneinasampleoffood,drinkingwater,
equipment,etc.Watersamplescanbefiltratedthroughanextremelyfinefilter.Thisfilteristhenplacedina
nutrientmedium.Microorganismsonthefilterthengrowtoformavisiblecolony.Harmfulmicroorganisms
canbedetectedinfoodbyplacingasampleinanutrientbrothdesignedtoenrichtheorganismsinquestion.
Variousmethods,suchasselectivemediaorpolymerasechainreaction,canthenbeusedfordetection.The
hygieneofhardsurfaces,suchascookingpots,canbetestedbytouchingthemwithasolidpieceofnutrient
mediumandthenallowingthemicroorganismstogrowonit.
Therearenoconditionswhereallmicroorganismswouldgrow,andthereforeoftenseveralmethodsare
needed.Forexample,afoodsamplemightbeanalyzedonthreedifferentnutrientmediumsdesignedto
indicatethepresenceof"total"bacteria(conditionswheremany,butnotall,bacteriagrow),molds
(conditionswherethegrowthofbacteriaispreventedby,e.g.,antibiotics)andcoliformbacteria(these
indicateasewagecontamination).
Seealso
Bacterium
Biologicalwarfare
Culturecollection
Cyanobacteria
Fungi
Helminths
Microbialbiogeography
Microbialintelligence
Nanobacterium
Nyloneatingbacteria
Petridish
Prokaryote
Protozoa
Soilcontamination
Staining
Virus
12/21
25/03/2015
References
1. MadiganM,MartinkoJ(editors)(2006).BrockBiologyofMicroorganisms(13thed.).PearsonEducation.
p.1096.ISBN032173551X.
2. RybickiEP(1990)."Theclassificationoforganismsattheedgeoflife,orproblemswithvirussystematics".S
AftJSci86:1826.ISSN00382353(https://www.worldcat.org/issn/00382353).
3. LWOFFA(1956)."Theconceptofvirus".J.Gen.Microbiol.17(2):23953.doi:10.1099/00221287172239
(https://dx.doi.org/10.1099%2F00221287172239).PMID13481308
(https://www.ncbi.nlm.nih.gov/pubmed/13481308).
4. UniversityofGeorgia(25August1998)."FirstEverScientificEstimateOfTotalBacteriaOnEarthShowsFar
GreaterNumbersThanEverKnownBefore"(http://www.sciencedaily.com/releases/1998/08/980825080732.htm).
ScienceDaily.Retrieved10November2014.
5. Zhang,K.DoseA.BiegerDose,R.Dillmann,M.Gill,O.Kerz,A.Klein,H.Meinert,T.Nawroth,S.Risi,C.
Stride(1995)."ERAexperiment"spacebiochemistry" ".AdvancesinSpaceResearch16(8):119129.
doi:10.1016/02731177(95)00280R(https://dx.doi.org/10.1016%2F02731177%2895%2900280R).
PMID11542696(https://www.ncbi.nlm.nih.gov/pubmed/11542696).
6. Vaisberg,HorneckGEschweilerU,ReitzG,WehnerJ,WillimekR,StrauchK.(1995)."Biologicalresponses
tospace:resultsoftheexperiment"ExobiologicalUnit"ofERAonEURECAI".AdvSpaceRes.16(8):10518.
Bibcode:1995AdSpR..16..105V(http://adsabs.harvard.edu/abs/1995AdSpR..16..105V).doi:10.1016/0273
1177(95)00279N(https://dx.doi.org/10.1016%2F02731177%2895%2900279N).PMID11542695
7. Staff(2014)."TheBiosphere"(http://www.agci.org/classroom/biosphere/index.php).AspenGlobalChange
Institute.Retrieved10November2014.
8. Choi,CharlesQ.(17March2013)."MicrobesThriveinDeepestSpotonEarth"
(http://www.livescience.com/27954microbesmarianatrench.html).LiveScience.Retrieved17March2013.
9. Glud,RonnieWenzhfer,FrankMiddelboe,MathiasOguri,KazumasaTurnewitsch,RobertCanfield,Donald
E.Kitazato,Hiroshi(17March2013)."Highratesofmicrobialcarbonturnoverinsedimentsinthedeepest
oceanictrenchonEarth"(http://www.nature.com/ngeo/journal/vaop/ncurrent/full/ngeo1773.html).Nature
Geoscience6(4):284.Bibcode:2013NatGe...6..284G(http://adsabs.harvard.edu/abs/2013NatGe...6..284G).
doi:10.1038/ngeo1773(https://dx.doi.org/10.1038%2Fngeo1773).Retrieved17March2013.
10. Oskin,Becky(14March2013)."Intraterrestrials:LifeThrivesinOceanFloor"
(http://www.livescience.com/27899oceansubsurfaceecosystemfound.html).LiveScience.Retrieved17March
2013.
11. Morelle,Rebecca(15December2014)."Microbesdiscoveredbydeepestmarinedrillanalysed"
(http://www.bbc.com/news/scienceenvironment30489814).BBCNews.Retrieved15December2014.
12. Fox,Douglas(20August2014)."Lakesundertheice:Antarcticassecretgarden"
(http://www.nature.com/news/lakesundertheiceantarcticassecretgarden1.15729).Nature(journal)512:
244246.doi:10.1038/512244a(https://dx.doi.org/10.1038%2F512244a).Retrieved21August2014.
13/21
25/03/2015
13. Mack,Eric(20August2014)."LifeConfirmedUnderAntarcticIceIsSpaceNext?"
(http://www.forbes.com/sites/ericmack/2014/08/20/lifeconfirmedunderantarcticiceisspacenext/).Forbes.
Retrieved21August2014.
14. ChristnerBC,MorrisCE,ForemanCM,CaiR,SandsDC(2008)."Ubiquityofbiologicalicenucleatorsin
snowfall".Science319(5867):1214.Bibcode:2008Sci...319.1214C
(http://adsabs.harvard.edu/abs/2008Sci...319.1214C).doi:10.1126/science.1149757
(https://dx.doi.org/10.1126%2Fscience.1149757).PMID18309078
15. 2002WHOmortalitydata(http://www.who.int/healthinfo/bodgbd2002revised/en/index.html)Accessed20
January2007
16. SchopfJ(2006)."FossilevidenceofArchaeanlife"
(http://www.journals.royalsoc.ac.uk/content/g38537726r273422/fulltext.pdf).PhilosTransRSocLondBBiolSci
361(1470):86985.doi:10.1098/rstb.2006.1834(https://dx.doi.org/10.1098%2Frstb.2006.1834).PMC1578735
(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1578735).PMID16754604
17. AltermannW,KazmierczakJ(2003)."Archeanmicrofossils:areappraisalofearlylifeonEarth".ResMicrobiol
154(9):6117.doi:10.1016/j.resmic.2003.08.006(https://dx.doi.org/10.1016%2Fj.resmic.2003.08.006).
18. CavalierSmithT(2006)."CellevolutionandEarthhistory:stasisandrevolution"
(http://www.journals.royalsoc.ac.uk/content/0164755512w92302/fulltext.pdf).PhilosTransRSocLondBBiol
Sci361(1470):9691006.doi:10.1098/rstb.2006.1842(https://dx.doi.org/10.1098%2Frstb.2006.1842).
PMC1578732(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1578732).PMID16754610
19. SchopfJ(1994)."Disparaterates,differingfates:tempoandmodeofevolutionchangedfromthePrecambrianto
thePhanerozoic"(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC44277).ProcNatlAcadSciUSA91(15):
673542.Bibcode:1994PNAS...91.6735S(http://adsabs.harvard.edu/abs/1994PNAS...91.6735S).
doi:10.1073/pnas.91.15.6735(https://dx.doi.org/10.1073%2Fpnas.91.15.6735).PMC44277
20. StanleyS(May1973)."AnEcologicalTheoryfortheSuddenOriginofMulticellularLifeintheLate
Precambrian"(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC433525).ProcNatlAcadSciUSA70(5):1486
21. DeLongE,PaceN(2001)."Environmentaldiversityofbacteriaandarchaea".SystBiol50(4):4708.
doi:10.1080/106351501750435040(https://dx.doi.org/10.1080%2F106351501750435040).PMID12116647
22. SchmidtA,RagazziE,CoppellottiO,RoghiG(2006)."AmicroworldinTriassicamber".Nature444(7121):
835.Bibcode:2006Natur.444..835S(http://adsabs.harvard.edu/abs/2006Natur.444..835S).doi:10.1038/444835a
(https://dx.doi.org/10.1038%2F444835a).PMID17167469(https://www.ncbi.nlm.nih.gov/pubmed/17167469).
14/21
25/03/2015
23. Schirber,Michael(July27,2014)."Microbe'sInnovationMayHaveStartedLargestExtinctionEventonEarth"
(http://www.space.com/26654microbeinnovationstartedlargestearthextinction.html).Space.com.Astrobiology
Magazine."....Thatspikeinnickelallowedmethanogenstotakeoff."
24. WolskaK(2003)."HorizontalDNAtransferbetweenbacteriaintheenvironment".ActaMicrobiolPol52(3):
23343.PMID14743976(https://www.ncbi.nlm.nih.gov/pubmed/14743976).
25. EnrightM,RobinsonD,RandleG,FeilE,GrundmannH,SprattB(May2002)."Theevolutionaryhistoryof
methicillinresistantStaphylococcusaureus(MRSA)"(http://www.pnas.org/cgi/pmidlookup?
view=long&pmid=12032344).ProcNatlAcadSciUSA99(11):768792.Bibcode:2002PNAS...99.7687E
(http://adsabs.harvard.edu/abs/2002PNAS...99.7687E).doi:10.1073/pnas.122108599
(https://dx.doi.org/10.1073%2Fpnas.122108599).PMC124322
26. Mahaviraisdated599BCE527BCE.See.Dundas,PaulJohnHinnelsed.(2002).TheJains.London:
Routledge.ISBN0415266068.p.24
27. Dundas,Paul(2002)p.88
28. Jaini,Padmanabh(1998).TheJainaPathofPurification.NewDelhi:MotilalBanarsidass.ISBN8120815785.
p.109
29. VarroOnAgriculture1,xiiLoeb
30. Tschanz,DavidW."ArabRootsofEuropeanMedicine"(http://www.hmc.org.qa/hmc/heartviews/hv
v4%20n2/9.htm).HeartViews4(2).
31. Colgan,Richard(2009).AdvicetotheYoungPhysician:OntheArtofMedicine(http://books.google.com/?
id=DoMVs4HuDAoC&pg=PA33&dq=avicenna+infectious+diseases+quarantine&q=avicenna%20infectious).
Springer.p.33.ISBN9781441910332.
32. Payne,A.S.TheCleereObserver:ABiographyofAntoniVanLeeuwenhoek,p.13,Macmillan,1970
33. LeeuwenhoekA(1753)."PartofaLetterfromMrAntonyvanLeeuwenhoek,concerningtheWormsinSheeps
Livers,Gnats,andAnimalculaintheExcrementsofFrogs"(http://www.journals.royalsoc.ac.uk/link.asp?
id=4j53731651310230).PhilosophicalTransactions(16831775)22(260276):50918.
doi:10.1098/rstl.1700.0013(https://dx.doi.org/10.1098%2Frstl.1700.0013).Retrieved30November2006.
34. LeeuwenhoekA(1753)."PartofaLetterfromMrAntonyvanLeeuwenhoek,F.R.S.concerningGreenWeeds
GrowinginWater,andSomeAnimalculaFoundaboutThem"(http://www.journals.royalsoc.ac.uk/link.asp?
id=fl73121jk4150280).PhilosophicalTransactions(16831775)23(277288):130411.
doi:10.1098/rstl.1702.0042(https://dx.doi.org/10.1098%2Frstl.1702.0042).Retrieved30November2006.
35. TheNobelPrizeinPhysiologyorMedicine1905(http://nobelprize.org/nobel_prizes/medicine/laureates/1905/)
Nobelprize.orgAccessed22November2006.
36. O'BrienS,GoedertJ(1996)."HIVcausesAIDS:Koch'spostulatesfulfilled".CurrOpinImmunol8(5):61318.
doi:10.1016/S09527915(96)800756(https://dx.doi.org/10.1016%2FS09527915%2896%29800756).
37. Borenstein,Seth(13November2013)."Oldestfossilfound:Meetyourmicrobialmom"
(http://apnews.excite.com/article/20131113/DAA1VSC01.html).APNews.Retrieved15November2013.
15/21
25/03/2015
38. Noffke,NoraChristian,ChristianWacey,DavidHazen,RobertM.(8November2013)."MicrobiallyInduced
SedimentaryStructuresRecordinganAncientEcosystemintheca.3.48BillionYearOldDresserFormation,
Pilbara,WesternAustralia"(http://online.liebertpub.com/doi/abs/10.1089/ast.2013.1030).Astrobiology(journal)
13(12):110324.doi:10.1089/ast.2013.1030(https://dx.doi.org/10.1089%2Fast.2013.1030).PMC3870916
(https://www.ncbi.nlm.nih.gov/pubmed/24205812).Retrieved15November2013.
39. CiccarelliFD,DoerksT,vonMeringC,CreeveyCJ,SnelB,BorkP(2006)."Towardautomaticreconstruction
ofahighlyresolvedtreeoflife".Science311(5765):12837.Bibcode:2006Sci...311.1283C
(http://adsabs.harvard.edu/abs/2006Sci...311.1283C).doi:10.1126/science.1123061
(https://dx.doi.org/10.1126%2Fscience.1123061).PMID16513982
40. GoldT(1992)."Thedeep,hotbiosphere"(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC49434).Proc.Natl.
Acad.Sci.U.S.A.89(13):60459.Bibcode:1992PNAS...89.6045G
(http://adsabs.harvard.edu/abs/1992PNAS...89.6045G).doi:10.1073/pnas.89.13.6045
(https://dx.doi.org/10.1073%2Fpnas.89.13.6045).PMC49434
41. WhitmanW,ColemanD,WiebeW(1998)."Prokaryotes:Theunseenmajority"
(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC33863).ProcNatlAcadSciUSA95(12):657883.
Bibcode:1998PNAS...95.6578W(http://adsabs.harvard.edu/abs/1998PNAS...95.6578W).
42. SchulzH,JorgensenB(2001)."Bigbacteria".AnnuRevMicrobiol55:10537.
doi:10.1146/annurev.micro.55.1.105(https://dx.doi.org/10.1146%2Fannurev.micro.55.1.105).PMID11544351
43. ShapiroJA(1998)."Thinkingaboutbacterialpopulationsasmulticellularorganisms"
(http://www.sci.uidaho.edu/newton/math501/Sp05/Shapiro.pdf).Annu.Rev.Microbiol.52:81104.
doi:10.1146/annurev.micro.52.1.81(https://dx.doi.org/10.1146%2Fannurev.micro.52.1.81).PMID9891794
44. JohnsborgO,EldholmV,HvarsteinLS(December2007)."Naturalgenetictransformation:prevalence,
mechanismsandfunction".Res.Microbiol.158(10):76778.doi:10.1016/j.resmic.2007.09.004
(https://dx.doi.org/10.1016%2Fj.resmic.2007.09.004).PMID17997281
45. SeealsoTransformation(genetics)
46. BernsteinH,BernsteinC,MichodRE(2012).DNArepairastheprimaryadaptivefunctionofsexinbacteriaand
eukaryotes.Chapter1:pp.149in:DNARepair:NewResearch,SakuraKimuraandSoraShimizueditors.Nova
Sci.Publ.,Hauppauge,N.Y.ISBN9781621008088
https://www.novapublishers.com/catalog/product_info.php?products_id=31918
47. SeealsoNaturalcompetence
16/21
25/03/2015
48. EagonR(1962)."PSEUDOMONASNATRIEGENS,AMARINEBACTERIUMWITHAGENERATION
TIMEOFLESSTHAN10MINUTES"(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC279347).JBacteriol
83(4):7367.PMC279347(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC279347).PMID13888946
49. WoeseC,KandlerO,WheelisM(1990)."Towardsanaturalsystemoforganisms:proposalforthedomains
Archaea,Bacteria,andEucarya"(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC54159).ProcNatlAcadSci
USA87(12):45769.Bibcode:1990PNAS...87.4576W(http://adsabs.harvard.edu/abs/1990PNAS...87.4576W).
50. DeRosaM,GambacortaA,GliozziA(1March1986)."Structure,biosynthesis,andphysicochemicalproperties
ofarchaebacteriallipids"(http://mmbr.asm.org/cgi/pmidlookup?view=long&pmid=3083222).Microbiol.Rev.50
(1):7080.PMC373054(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC373054).PMID3083222
51. RobertsonC,HarrisJ,SpearJ,PaceN(2005)."PhylogeneticdiversityandecologyofenvironmentalArchaea".
CurrOpinMicrobiol8(6):63842.doi:10.1016/j.mib.2005.10.003
(https://dx.doi.org/10.1016%2Fj.mib.2005.10.003).PMID16236543
52. KarnerMB,DeLongEF,KarlDM(2001)."ArchaealdominanceinthemesopelagiczoneofthePacificOcean".
Nature409(6819):50710.doi:10.1038/35054051(https://dx.doi.org/10.1038%2F35054051).PMID11206545
53. SinningheDamstJS,RijpstraWI,HopmansEC,PrahlFG,WakehamSG,SchoutenS(June2002).
"DistributionofMembraneLipidsofPlanktonicCrenarchaeotaintheArabianSea"
(http://aem.asm.org/cgi/pmidlookup?view=long&pmid=12039760).Appl.Environ.Microbiol.68(6):29973002.
doi:10.1128/AEM.68.6.29973002.2002(https://dx.doi.org/10.1128%2FAEM.68.6.29973002.2002).
54. LeiningerS,UrichT,SchloterMetal.(2006)."Archaeapredominateamongammoniaoxidizingprokaryotesin
soils".Nature442(7104):8069.Bibcode:2006Natur.442..806L
(http://adsabs.harvard.edu/abs/2006Natur.442..806L).doi:10.1038/nature04983
(https://dx.doi.org/10.1038%2Fnature04983).PMID16915287
55. Eukaryota:MoreonMorphology.(http://www.ucmp.berkeley.edu/alllife/eukaryotamm.html)(Retrieved10
October2006)
56. DyallS,BrownM,JohnsonP(2004)."Ancientinvasions:fromendosymbiontstoorganelles".Science304
(5668):2537.Bibcode:2004Sci...304..253D(http://adsabs.harvard.edu/abs/2004Sci...304..253D).
doi:10.1126/science.1094884(https://dx.doi.org/10.1126%2Fscience.1094884).PMID15073369
57. Seecoenocyte.
58. AlsoseeMeiosis.
17/21
25/03/2015
59. CavalierSmithT(1December1993)."Kingdomprotozoaandits18phyla"
(http://mmbr.asm.org/cgi/pmidlookup?view=long&pmid=8302218).Microbiol.Rev.57(4):95394.
60. CorlissJO(1992)."Shouldtherebeaseparatecodeofnomenclaturefortheprotists?".BioSystems28(13):1
14.doi:10.1016/03032647(92)90003H(https://dx.doi.org/10.1016%2F03032647%2892%2990003H).
61. DevreotesP(1989)."Dictyosteliumdiscoideum:amodelsystemforcellcellinteractionsindevelopment".
Science245(4922):10548.Bibcode:1989Sci...245.1054D(http://adsabs.harvard.edu/abs/1989Sci...245.1054D).
doi:10.1126/science.2672337(https://dx.doi.org/10.1126%2Fscience.2672337).PMID2672337
62. SlapetaJ,MoreiraD,LpezGarcaP(2005)."Theextentofprotistdiversity:insightsfrommolecularecologyof
freshwatereukaryotes"(http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1559898/).Proc.Biol.Sci.272(1576):
207381.doi:10.1098/rspb.2005.3195(https://dx.doi.org/10.1098%2Frspb.2005.3195).PMC1559898
63. MoreiraD,LpezGarcaP(2002)."Themolecularecologyofmicrobialeukaryotesunveilsahiddenworld"
(http://download.bioon.com.cn/view/upload/month_0803/20080326_daa08a6fdb5d38e3a0d8VBrocN3WtOdR.attac
h.pdf).TrendsMicrobiol.10(1):318.doi:10.1016/S0966842X(01)022570
(https://dx.doi.org/10.1016%2FS0966842X%2801%29022570).PMID11755083
64. LapinskiJ,TunnacliffeA(2003)."Anhydrobiosiswithouttrehaloseinbdelloidrotifers".FEBSLett.553(3):
38790.doi:10.1016/S00145793(03)010627(https://dx.doi.org/10.1016%2FS00145793%2803%29010627).
65. KumamotoCA,VincesMD(2005)."ContributionsofhyphaeandhyphacoregulatedgenestoCandidaalbicans
virulence".Cell.Microbiol.7(11):154654.doi:10.1111/j.14625822.2005.00616.x
(https://dx.doi.org/10.1111%2Fj.14625822.2005.00616.x).PMID16207242
66. Thomas,DavidC.(2002).Seaweeds.London:NaturalHistoryMuseum.ISBN0565091751.
67. SzewzykU,SzewzykR,StenstrmT(1994)."Thermophilic,anaerobicbacteriaisolatedfromadeepboreholein
graniteinSweden"(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC43253).ProcNatlAcadSciUSA91(5):
68. HorneckG(1981)."Survivalofmicroorganismsinspace:areview".AdvSpaceRes1(14):3948.
doi:10.1016/02731177(81)902416(https://dx.doi.org/10.1016%2F02731177%2881%29902416).
69. Strain121,ahyperthermophilicarchaea,hasbeenshowntoreproduceat121C(250F),andsurviveat130C
(266F).[1](http://www.nsf.gov/od/lpa/news/03/pr0384.htm)
18/21
25/03/2015
70. SomePsychrophilicbacteriacangrowat17C(1F),[2](http://news.bbc.co.uk/1/hi/sci/tech/827063.stm)and
cansurvivenearabsolutezero.[3](http://science.nasa.gov/newhome/headlines/ast01sep98_1.htm)
71. PicrophiluscangrowatpH0.06.[4](http://www.rcn.montana.edu/resources/organisms/organisminfo.aspx?
nav=11&tid=1298&did=1&nid=82076&lid=9)
72. ThealkaliphilicbacteriaBacillusalcalophiluscangrowatuptopH11.5.[5]
(http://jb.asm.org/cgi/reprint/185/2/461.pdf)
73. DyallSmith,Mike,HALOARCHAEA(http://www.microbiol.unimelb.edu.au/people/dyallsmith/),Universityof
Melbourne.SeealsoHaloarchaea.
74. ThepiezophilicbacteriaHalomonassalariarequiresapressureof1,000atmnanobes,aspeculativeorganism,
havebeenreportedlyfoundintheearth'scrustat2,000atm.[6](http://www.microscopyuk.org.uk/index.html?
http://www.microscopyuk.org.uk/nanobes/nanopaper.html)
75. SeeDeinococcusradiodurans
76. CavicchioliR(2002)."Extremophilesandthesearchforextraterrestriallife".Astrobiology2(3):28192.
Bibcode:2002AsBio...2..281C(http://adsabs.harvard.edu/abs/2002AsBio...2..281C).
doi:10.1089/153110702762027862(https://dx.doi.org/10.1089%2F153110702762027862).PMID12530238
77. BareaJ,PozoM,AzcnR,AzcnAguilarC(2005)."Microbialcooperationintherhizosphere".JExpBot56
(417):176178.doi:10.1093/jxb/eri197(https://dx.doi.org/10.1093%2Fjxb%2Feri197).PMID15911555
78. Gillen,AlanL.(2007).TheGenesisofGerms:TheOriginofDiseasesandtheComingPlagues.NewLeaf
PublishingGroup.p.10.ISBN0890514933.
79. "DairyMicrobiology"(http://www.foodsci.uoguelph.ca/dairyedu/micro.html).UniversityofGuelph.Retrieved
9October2006.
80. Gray,N.F.(2004).BiologyofWastewaterTreatment.ImperialCollegePress.p.1164.ISBN1860943322.
81. Kitani,OsumuandCarlW.Hall(1989).BiomassHandbook.Taylor&FrancisUS.p.256.ISBN288124269
3.
82. Pimental,David(2007).Food,Energy,andSociety.CRCPress.p.289.ISBN1420046675.
83. Tickell,Joshuaetal.(2000).FromtheFryertotheFuelTank:TheCompleteGuidetoUsingVegetableOilasan
AlternativeFuel.BiodieselAmerica.p.53.ISBN0970722702.
84. Inslee,Jayetal.(2008).Apollo'sFire:IgnitingAmerica'sCleanEnergyEconomy.IslandPress.p.157.ISBN1
597261750.
85. BiologytextbookforclassXII.Nationalcouncilofeducationalresearchandtraining.p.183.ISBN817450639
X.
86. CastrilloJI,OliverSG(2004)."Yeastasatouchstoneinpostgenomicresearch:strategiesforintegrative
analysisinfunctionalgenomics"(http://www.jbmb.or.kr/fulltext/jbmb/view.php?vol=37&page=93).J.Biochem.
Mol.Biol.37(1):93106.doi:10.5483/BMBRep.2004.37.1.093
(https://dx.doi.org/10.5483%2FBMBRep.2004.37.1.093).PMID14761307
87. SuterB,AuerbachD,StagljarI(2006)."Yeastbasedfunctionalgenomicsandproteomicstechnologies:thefirst
15yearsandbeyond".BioTechniques40(5):62544.doi:10.2144/000112151
(https://dx.doi.org/10.2144%2F000112151).PMID16708762(https://www.ncbi.nlm.nih.gov/pubmed/16708762).
19/21
25/03/2015
88. SunnerhagenP(2002)."ProspectsforfunctionalgenomicsinSchizosaccharomycespombe".Curr.Genet.42(2):
7384.doi:10.1007/s0029400203356(https://dx.doi.org/10.1007%2Fs0029400203356).PMID12478386
89. Soni,S.K.(2007).Microbes:ASourceofEnergyfor21stCentury.NewIndiaPublishing.ISBN8189422146.
90. Moses,Vivianetal.(1999).Biotechnology:TheScienceandtheBusiness.CRCPress.p.563.ISBN905702
4071.
91. Langford,RolandE.(2004).IntroductiontoWeaponsofMassDestruction:Radiological,Chemical,and
Biological.WileyIEEE.p.140.ISBN0471465607.
92. O'HaraA,ShanahanF(2006)."Thegutfloraasaforgottenorgan"
(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1500832).EMBORep7(7):68893.
doi:10.1038/sj.embor.7400731(https://dx.doi.org/10.1038%2Fsj.embor.7400731).PMC1500832
93. EckburgP,LeppP,RelmanD(2003)."ArchaeaandTheirPotentialRoleinHumanDisease"
(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC145348).InfectImmun71(2):5916.
doi:10.1128/IAI.71.2.591596.2003(https://dx.doi.org/10.1128%2FIAI.71.2.591596.2003).PMC145348
94. LeppP,BrinigM,OuverneyC,PalmK,ArmitageG,RelmanD(2004)."MethanogenicArchaeaandhuman
periodontaldisease"(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC395942).ProcNatlAcadSciUSA101
(16):617681.Bibcode:2004PNAS..101.6176L(http://adsabs.harvard.edu/abs/2004PNAS..101.6176L).
doi:10.1073/pnas.0308766101(https://dx.doi.org/10.1073%2Fpnas.0308766101).PMC395942
Externallinks
OurMicrobialPlanet(http://dels.nas.edu/metagenomics)AfreeposterfromtheNationalAcademyof
Sciencesaboutthepositiverolesofmicroorganisms.
"UnchartedMicrobialWorld:MicrobesandTheirActivitiesintheEnvironment"
(http://www.asm.org/ASM/files/ccLibraryFiles/Filename/000000003691/Uncharted_Microbial_Worl
d.pdf)ReportfromtheAmericanAcademyofMicrobiology
UnderstandingOurMicrobialPlanet:TheNewScienceofMetagenomics
(http://dels.nas.edu/dels/rpt_briefs/metagenomics_final.pdf)A20pageeducationalbookletproviding
abasicoverviewofmetagenomicsandourmicrobialplanet.
TreeofLifeEukaryotes(http://tolweb.org/Eukaryotes/3)
MicrobeNewsfromGenomeNewsNetwork
(http://www.genomenewsnetwork.org/categories/index/microbes.php)
MedicalMicrobiology(http://gsbs.utmb.edu/microbook/toc.htm)Onlinetextbook
20/21
25/03/2015
Throughthemicroscope:Alookatallthingssmall(http://www.microbiologytext.com/index.php?
module=Book&func=toc&book_id=4)OnlinemicrobiologytextbookbyTimothyPaustianandGary
Roberts,UniversityofWisconsinMadison
Microorganismsinthepondwater(https://www.youtube.com/watch?v=sDacX2Xs0X4)onYouTube
Methanespewingmicrobeblamedinworstmassextinction.CBCNews
(http://www.cbc.ca/news/technology/methanespewingmicrobeblamedinworstmassextinction
1.2595797)
Retrievedfrom"http://en.wikipedia.org/w/index.php?title=Microorganism&oldid=652763264"
Categories: Microorganisms
Thispagewaslastmodifiedon20March2015,at18:03.
TextisavailableundertheCreativeCommonsAttributionShareAlikeLicenseadditionaltermsmay
apply.Byusingthissite,youagreetotheTermsofUseandPrivacyPolicy.Wikipediaisa
registeredtrademarkoftheWikimediaFoundation,Inc.,anonprofitorganization.
21/21

Microorganism

Diunggah oleh

Informasi Dokumen

Hak Cipta

Format Tersedia

Bagikan dokumen Ini

Bagikan atau Tanam Dokumen

Opsi Berbagi

Apakah menurut Anda dokumen ini bermanfaat?

Apakah konten ini tidak pantas?

Hak Cipta:

Format Tersedia

Microorganism

Diunggah oleh

Hak Cipta:

Format Tersedia

25/03/2015

Anda mungkin juga menyukai