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OperonsandProkaryoticGeneRegulation

By:AmyRalston,Ph.D.(WriteScienceRight)©2008NatureEducation

Citation:Ralston,A.(2008)Operonsandprokaryoticgeneregulation.Nature

Education1(1):216

Howdobacteriaadaptsoquicklytotheirenvironments?Partoftheanswertothisquestion

liesinclustersofcoregulatedgenescalledoperons.

Bacteriaaretypicallyexposedtoanever­changingenvironmentinwhichnutrientavailabilitymayincreaseor

decreaseradically.Bacteriarespondtosuchvariationsintheirenvironmentbyalteringtheirgeneexpression

pattern;thus,theyexpressdifferentenzymesdependingonthecarbonsourcesandothernutrientsavailableto

them.Itwouldbewastefultosynthesize,forexample,lactose­metabolizingenzymesintheabsenceoflactose.

However,whenlactoseistheonlyavailablecarbonsource,bacteriamustquicklyinducelactose­metabolizing

enzymes,orelsetheywilldie.Inbacteria,thissortofgeneticregulationismediatedattheleveloftranscription.

BacterialOperonsAreCoregulatedGeneClusters

Bacterialgenesareorganizedintooperons,orclustersofcoregulatedgenes.Inadditiontobeingphysically

closeinthegenome,thesegenesareregulatedsuchthattheyareallturnedonorofftogether.Groupingrelated

genesunderacommoncontrolmechanismallowsbacteriatorapidlyadapttochangesintheenvironment.

Thebest­studiedexamplesofoperonsarefromthebacteriumEscherichiacoli(E.coli),andtheyinvolvetheenzymes

oflactosemetabolismandtryptophanbiosynthesis.Becausethelactose(lac)operonsharesmanyfeatureswith

otheroperons,itsorganizationandregulationaredescribedindetailbelow.

ThelacOperon

NobellaureatesFrancoisJacobandJacquesMonod conductedmuchoftheseminalworkdescribingthelac operon'sstructureandcontrolmechanisms.By examiningmutantstrainsofE.colithatexhibited defectsinlactosemetabolism,JacobandMonodwere abletolearnhowthelacoperonisregulatedto

metabolizelactose(Jacob&Monod,1962).Theduo

notedthatthelacoperoncontainsthreegenesthat

encodeproteinsinvolvedinlactosemetabolism.These

arereferredtoaslacz,lacy,andlaca.Thelaczgene

encodesbeta­galactosidase,thelacygeneencodesa

permease,andthelacageneencodesthe

transacetylaseenzyme.Together,thesegeneproducts

acttoimportlactoseintocellsandbreakitdownforuse

asafoodsource.Asinotheroperons,thegenesofthe

lacoperonliealongacontiguousstretchofDNAsuch

thattheirexpressioncanbeeasilycoregulated.In

additiontotheseso­calledstructuralgenes,thelac

operonalsocontainsothersequencesthatdirectthe

bacterialgeneexpressionmachinery.

bacterial geneexpression machinery. Figure1:Thelacoperonin E.coli . Threelactosemetabolismgenes(

Figure1:ThelacoperoninE.coli . E.coli.

Threelactosemetabolismgenes(lacZ,lacY,and

lacA)areorganizedtogetherinaclustercalledthe

lacoperon.Thecoordinatedtranscriptionand

translationofthelacoperonstructuralgenesis

supportedbyasharedpromoter,operator,and

terminator.Alacregulatorgenewithitspromoter

isfoundjustoutsidethelacoperon.

©2013NatureEducationAdaptedfromPierce,Benjamin.

Genetics:AConceptualApproach,2nded.Allrightsreserved.

Genetics:AConceptualApproach ,2nded.Allrightsreserved. Theorganizationofgenesintoanoperonallowsfor

Theorganizationofgenesintoanoperonallowsfor

simultaneousexpressionofallthegenesthatare

locatedincis(i.e.,onthesamecontiguouspieceof

DNA)intheoperon.Severalfeaturescontributetothis

characteristicofoperons(Figure1).First,allofthe

operon'sgenesaredownstreamofasinglepromoter.Thispromoterservesasarecognitionsiteforthe

transcriptionalmachineryoftheRNApolymerasecomplex.Second,allgenesinanoperonactuallybecomepart

ofasinglemessengerRNAmolecule,whichissubsequentlytranslatedintoindividualproteingeneproducts.

Severalotherregulatorysequencesalsoensurecoordinatedregulationofthelacoperon.Theseincludethe

operatorandtheterminator.TheoperatorisaspecialDNAsequencelocatedbetweenthepromotersequence

andthestructuralgenesthatenablesrepressionoftheentirelacoperon,followingbindingbytheinhibitor(laci)

protein.Expressionofthelacoperonis,infact,regulatedbythepresenceoflactoseitself.Theabilitytoturn

lactose­metabolizinggenesonoroffasagroupthereforeprovidesanefficientwaytoquicklyadaptto

environmentalchanges.Theterminator,ontheotherhand,instructsthetranscriptionalmachinerytoterminate

transcription.Assuch,thepromoterservesasatranscriptionalstartsite,theterminatorservesasastopsite,

andtheoperatorhelpsdeterminewhethertranscriptionwilloccur.

Becauseofthecommoncontrolmechanismforallofthegenesinthelacoperon,mutationsinthisoperoncan

havemultiple,orpleiotropic,effects.Forexample,mutationsaffectingthepromotercanpreventallofthe

operon'sgenesfrombeingexpressed,becauseRNApolymerasewillbeunabletobindandcommence

transcription.Othermutationsmayaffectexpressionofonlysomeofthegenesintheoperon.Forinstance,

nonsensemutationsinthelaczgeneencodingbeta­galactosidasemightinterferewithexpressionofdownstream

permeaseandtransacetylasegenesbycausingtheribosometofalloffpriortotheirtranslation.Thiskindof

mutationissaidtohavea"polar"effectonthepathwayinthatitaffectsdownstreamgenesbutnotupstream

genes.

Thus,althoughbacteriamaybeconsideredsimplerorganismsthanhumans,itisclearthatbacterialgene

regulationisextremelyefficientandthatthebacterialgenomeishighlyorganized.Bacteriaappeartobe

perfectlyadaptedtoavarietyofenvironments,andtheyarereadytorespondtowhateverenvironmental

changestheyencounterbyemployingelegantandcomplexregulatorymechanisms.

ReferencesandRecommendedReading

Jacob,F.,&Monod,J.Ontheregulationofgeneactivity.ColdSpringHarborSymposiaonQuantitativeBiology26,193–211(1962)

Lawrence,J.G.Sharedstrategiesingeneorganizationamongprokaryotesandeukaryotes.Cell110,407–413(2002)