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PROCARYOTICLIFE
TheBacteriaareagroupofsingle-cellmicroorganismswithprocaryoticcellularconfiguration.Thegeneticmaterial(DNA)ofprocaryoticcellsexistsunboundinthecytoplasmofthecells.Thereisnonuclearmembrane,whichisthedefinitivecharacteristicofeucaryoticcellssuchasthosethatmakeup,fungi,protista,plantsandanimals.Untilrecently,bacteriaweretheonlyknowntypeofprocaryoticcell,andthedisciplineofbiologyrelatedtotheirstudyiscalledbacteriology.Inthe1980'
s,withtheoutbreakofmoleculartechniquesappliedtophylogenyoflife,anothergroupofprocaryoteswasdefinedandinformallynamed"
archaebacteria"
.ThisgroupofprocaryoteshassincebeenrenamedArchaeaandhasbeenawardedbiologicalDomainstatusonthelevelwithBacteriaandEucarya.Thecurrentscienceofbacteriologyincludesthestudyofbothdomainsofprocaryoticcells,butthename"
bacteriology"
isnotlikelytochangetoreflecttheinclusionofarchaeainthediscipline.Actually,manyarchaeahavebeenstudiedasintensivelyandaslongastheirbacterialcounterparts,exceptwiththenotionthattheywerebacteria.
Figure1.ThecyanobacteriumAnabaena.AmericanSocietyforMicrobiology.Two(notuncommon)exceptionsthatprocaryotesareunicellularandundifferentiatedareseeninAnabaena:
1.Theorganismlivesasamulticellularfilamentorchainofcells.Procaryotesareconsidered"
unicellularorganisms"
becauseallthecellsinafilamentorcolonyareofthesametype,andanyoneindividualcellcangiverisetoanexactfilamentorcolony;
2.Thepredominantphotosynthetic(brightyellow-green)cellsdodifferentiateintoanothertypeofcell:
theobviouslylarge"
empty"
cellsoccasionallyseenalongafilamentaredifferentiatedcellsinwhichnitrogenfixation,butnotphotosynthesis,takesplace.
TheOriginofLife
WhenlifearoseonEarthabout4billionyearsago,thefirsttypesofcellstoevolvewereprocaryoticcells.Forapproximately2billionyears,procaryotic-typecellsweretheonlyformoflifeonEarth.Theoldestknownsedimentaryrocks,fromGreenland,areabout3.8billionyearsold.Theoldestknownfossilsareprocaryoticcells,3.5billionyearsinage,foundinWesternAustraliaandSouthAfrica.Thenatureofthesefossils,andthechemicalcompositionoftherocksinwhichtheyarefound,indicatesthatlithotrophicandfermentativemodesofmetabolismwerethefirsttoevolveinearlyprocaryotes.Photosynthesisdevelopedinbacteriaabitlater,atleast3billionyearsago.Anoxygenicphotosynthesis(bacterialphotosynthesis,whichisanaerobicanddoesnotproduceO2)precededoxygenicphotosynthesis(plant-typephotosynthesis,whichyieldsO2).However,oxygenicphotosynthesisalsoaroseinprocaryotes,specificallyinthecyanobacteria,whichexistedmillionsofyearsbeforetheevolutionofgreenalgaeandplants.Larger,morecomplicatedeucaryoticcellsdidnotappearuntilmuchlater,between1.5and2billionyearsago.
Figure2.OpalescentPoolinYellowstoneNationalPark,WyomingUSA.K.Todar.ConditionsforlifeinthisenvironmentaresimilartoEarthover2billionyearsago.Inthesetypesofhotsprings,theorange,yellowandbrowncolorsareduetopigmentedphotosyntheticbacteriawhichmakeupthemicrobialmats.Thematsareliterallyteemingwithbacteria.SomeofthesebacteriasuchasSynechococcusconductoxygenicphotosynthesis,whileotherssuchasChloroflexusconductanoxygenicphotosynthesis.Othernon-photosyntheticbacteria,aswellasthermophilicandacidophilicArchaea,arealsoresidentsofthehotspringcommunity.
Thearchaeaandbacteriadifferfundamentallyintheirstructurefromeucaryoticcells,whichalwayscontainamembrane-enclosednucleus,multiplechromosomes,andvariousothermembranousorganellessuchasmitochondria,chloroplasts,thegolgiapparatus,vacuoles,etc.Unlikeplantsandanimals,archaeaandbacteriaareunicellularorganismsthatdonotdevelopordifferentiateintomulticellularforms.Somebacteriagrowinfilamentsormassesofcells,buteachcellinthecolonyisidenticalandcapableofindependentexistence.Thecellsmaybeadjacenttooneanotherbecausetheydidnotseparateaftercelldivisionorbecausetheyremainedenclosedinacommonsheathorslimesecretedbythecells,buttypicallythereisnocontinuityorcommunicationbetweenthecells.
TheUniversalTreeofLife
OnthebasisofsmallsubunitribosomalRNA(ssrRNA)analysis,thecontemporaryTreeofLifegivesrisetothreecellular"
Domains"
:
Archaea,Bacteria,andEucarya(Figure3).Bacteria(formerlyknownaseubacteria)andArchaea(formerlycalledarchaebacteria)sharetheprocaryotictypeofcellularconfiguration,butotherwisearenotrelatedtooneanotheranymorecloselythantheyaretotheeucaryoticdomain,Eucarya.Betweenthetwoprocaryotes,ArchaeaareapparentlymorecloselyrelatedtoEucaryathanaretheBacteria.Eucaryaconsistsofalleucaryoticcell-types,includingprotista,fungi,plantsandanimals.
Figure3.TheUniversalTreeofLifeasderivedfromsequencingofssrRNA.N.Pace.Notethethreemajordomainsoflivingorganisms:
Archaea,BacteriaandEucarya.The"
evolutionarydistance"
betweentwoorganismsisproportionaltothemeasurabledistancebetweentheendofabranchtoanodetotheendofacomparativebranch.
Forexample,inEucarya,humans(Homo)aremorecloselyrelatedtocorn(Zea)thantoslimemolds(Dictyostelium);
orinBacteria,E.coliismorecloselyrelatedtoAgrobacteriumthantoThermus.
SizeandDistributionofBacteriaandArchaea
Mostprocaryoticcellsareverysmallcomparedtoeucaryoticcells.Atypicalbacterialcellisabout1micrometerindiameterorwidth,whilemosteucaryoticcellsarefrom10to100micrometersindiameter.Eucaryoticcellshaveamuchgreatervolumeofcytoplasmandamuchlowersurface:
volumeratiothanprocaryoticcells.Atypicalprocaryoticcellisaboutthesizeofaeucaryoticmitochondrion.Sinceprocaryotesaretoosmalltobeseenexceptwiththeaidofamicroscope,itisusuallynotappreciatedthattheyarethemostabundantformoflifeontheplanet,bothintermsofbiomassandtotalnumbersofspecies.Forexample,inthesea,procaryotesmakeup90percentofthetotalcombinedweightofallorganisms.Inasinglegramoffertileagriculturalsoiltheremaybeinexcessof109bacterialcells,outnumberingalleucaryoticcellsthereby10,000:
1.About3,000distinctspeciesofbacteriaandarchaeaarerecognized,butthisnumberisprobablylessthanonepercentofallthespeciesinnature.Theseunknownprocaryotes,farinexcessofundiscoveredorunstudiedplants,areatremendousreserveofgeneticmaterialandgeneticinformationinnaturethatawaitsexploitation.
Procaryotesarefoundinallofthehabitatswhereeucaryoteslive,but,aswell,inmanynaturalenvironmentsconsideredtooextremeorinhospitableforeucaryoticcells.Thus,theouterlimitsoflifeonEarth(hottest,coldest,driest,etc.)areusuallydefinedbytheexistenceofprocaryotes.Whereeucaryotesandprocaryoteslivetogether,theremaybemutualisticassociationsbetweentheorganismsthatallowbothtosurviveorflourish.Theorganellesofeucaryotes(mitochondriaandchloroplasts)arethoughttoberemnantsofBacteriathatinvaded,orwerecapturedby,primitiveeucaryotesintheevolutionarypast.Numeroustypesofeucaryoticcellsthatexisttodayareinhabitedbyendosymbioticprocaryotes.
Fromametabolicstandpoint,theprocaryotesareextraordinarilydiverse,andtheyexhibitseveraltypesofmetabolismthatarerarelyorneverseenineucaryotes.Forexample,thebiologicalprocessesofnitrogenfixation(conversionofatmosphericnitrogengastoammonia)andmethanogenesis(productionofmethane)aremetabolically-uniquetoprocaryotesandhaveanenormousimpactonthenitrogenandcarboncyclesinnature.UniquemechanismsforenergyproductionandphotosynthesisarealsoseenamongtheArchaeaandBacteria.
Thelivesofplantsandanimalsaredependentupontheactivitiesofbacterialcells.Bacteriaandarchaeaenterintovarioustypesofsymbioticrelationshipswithplantsandanimalsthatusuallybenefitbothorganisms,althoughafewbacteriaareagentsofdisease.
Themetabolicactivitiesofprocaryotesinsoilhabitatshaveanenormousimpactonsoilfertilitythatcanaffectagriculturalpracticesandcropyields.Intheglobalenvironment,procaryotesareabsolutelyessentialtodrivethecyclesofelementsthatmakeuplivingsystems,i.e.,thecarbon,oxygen,nitrogenandsulfurcycles.Theoriginsoftheplantcellchloroplastandplant-type(oxygenic)photosynthesisarefoundinprocaryotes.Mostoftheearth'
satmosphericoxygenmayhavebeenproducedbyfree-livingbacterialcells.ThebacteriafixnitrogenandasubstantialamountofCO2,aswell.
Bacteriaorbacterialproducts(includingtheirgenes)canbeusedtoincreasecropyieldorplantresistancetodisease,ortocureorpreventplantdisease.Bacterialproductsincludeantibioticstofightinfectiousdisease,aswellascomponentsforvaccinesusedtopreventinfectiousdisease.Becauseoftheirsimplicityandourrelativeunderstandingoftheirbiologicalprocesses,thebacteriaprovideconvenientlaboratorymodelsforstudyofthemolecularbiology,genetics,andphysiologyofalltypesofcells,includingplantandanimalcells.
STRUCTUREANDFUNCTIONOFPROCARYOTICCELLS
Procaryoticcellshavethreearchitecturalregions(Figure4):
appendages(proteinsattachedtothecellsurface)intheformofflagellaandpili;
acellenvelopeconsistingofacapsule,cellwallandplasma