Caenorhabditis elegans - Wikipedia

Caenorhabditis elegans is a free-living transparent nematode about 1 mm in length that lives in temperate soil environments. It is the type species of its ... Caenorhabditiselegans FromWikipedia,thefreeencyclopedia Jumptonavigation Jumptosearch Free-livingspeciesofnematode "C.elegans"redirectshere.Forotheruses,seeC.elegans(disambiguation). Caenorhabditiselegans AnadulthermaphroditeC.elegansworm Scientificclassification Kingdom: Animalia Phylum: Nematoda Class: Chromadorea Order: Rhabditida Family: Rhabditidae Genus: Caenorhabditis Species: C. elegans Binomialname Caenorhabditiselegans(Maupas,1900)[1] Subspecies Caenorhabditiselegansvar.Bergerac[2](forinstancestrainBO)[3] Caenorhabditiselegansvar.Bristol[4](forinstancestrainN2)[5] Caenorhabditiselegans(/ˌsiːnoʊræbˈdaɪtəsˈɛləɡæns/[6])isafree-livingtransparentnematodeabout1 mminlength[7]thatlivesintemperatesoilenvironments.Itisthetypespeciesofitsgenus.[8]ThenameisablendoftheGreekcaeno-(recent),rhabditis(rod-like)[9]andLatinelegans(elegant).In1900,MaupasinitiallynameditRhabditideselegans.OscheplaceditinthesubgenusCaenorhabditisin1952,andin1955,DoughertyraisedCaenorhabditistothestatusofgenus.[10] C.elegansisanunsegmentedpseudocoelomateandlacksrespiratoryorcirculatorysystems.[11]Mostofthesenematodesarehermaphroditesandafewaremales.[12]Maleshavespecialisedtailsformatingthatincludespicules. In1963,SydneyBrennerproposedresearchintoC.elegans,primarilyintheareaofneuronaldevelopment.In1974,hebeganresearchintothemolecularanddevelopmentalbiologyofC.elegans,whichhassincebeenextensivelyusedasamodelorganism.[13]Itwasthefirstmulticellularorganismtohaveitswholegenomesequenced,andasof2019[update],istheonlyorganismtohaveitsconnectome(neuronal"wiringdiagram")completed.[14][15][16] Contents 1Anatomy 1.1Gutgranules 2Reproduction 2.1Sexdetermination 2.1.1Evolution 3Development 3.1Embryonicdevelopment 3.1.1Axisformation 3.1.2Gastrulation 3.2Post-embryonicdevelopment 4Ecology 5Useasamodelorganism 5.1Celllineagemapping 5.2Programmedcelldeath 5.3RNAinterferenceandgenesilencing 5.4Celldivisionandcellcycle 5.5Drugabuseandaddiction 5.6Biologicaldatabases 5.7Ageing 5.8Sleep 5.9Sensorybiology 5.10Spaceflightresearch 6Genetics 6.1Genome 6.1.1Sizeandgenecontent 6.1.2Relatedgenomes 6.2Othergeneticstudies 7Scientificcommunity 8Seealso 9References 10Furtherreading 11Externallinks Anatomy[edit] Movementofwild-typeC.elegans C.elegansisunsegmented,vermiform,andbilaterallysymmetrical.Ithasacuticle(atoughoutercovering,asanexoskeleton),fourmainepidermalcords,andafluid-filledpseudocoelom(bodycavity).Italsohassomeofthesameorgansystemsaslargeranimals.Aboutoneinathousandindividualsismaleandtherestarehermaphrodites.[17]ThebasicanatomyofC.elegansincludesamouth,pharynx,intestine,gonad,andcollagenouscuticle.Likeallnematodes,theyhaveneitheracirculatorynorarespiratorysystem.Thefourbandsofmusclesthatrunthelengthofthebodyareconnectedtoaneuralsystemthatallowsthemusclestomovetheanimal'sbodyonlyasdorsalbendingorventralbending,butnotleftorright,exceptforthehead,wherethefourmusclequadrantsarewiredindependentlyfromoneanother.Whenawaveofdorsal/ventralmusclecontractionsproceedsfromthebacktothefrontoftheanimal,theanimalispropelledbackwards.Whenawaveofcontractionsisinitiatedatthefrontandproceedsposteriorlyalongthebody,theanimalispropelledforwards.Becauseofthisdorsal/ventralbiasinbodybends,anynormalliving,movingindividualtendstolieoneitheritsleftsideoritsrightsidewhenobservedcrossingahorizontalsurface.Asetofridgesonthelateralsidesofthebodycuticle,thealae,isbelievedtogivetheanimaladdedtractionduringthesebendingmotions. VideoofC.eleganswithpencilforscale Inrelationtolipidmetabolism,C.elegansdoesnothaveanyspecializedadiposetissues,apancreas,aliver,orevenbloodtodelivernutrientscomparedtomammals.Neutrallipidsareinsteadstoredintheintestine,epidermis,andembryos.Theepidermiscorrespondstothemammalianadipocytesbybeingthemaintriglyceridedepot.[18] ThepharynxisamuscularfoodpumpintheheadofC.elegans,whichistriangularincross-section.Thisgrindsfoodandtransportsitdirectlytotheintestine.Asetof"valvecells"connectsthepharynxtotheintestine,buthowthisvalveoperatesisnotunderstood.Afterdigestion,thecontentsoftheintestinearereleasedviatherectum,asisthecasewithallothernematodes.[19]Nodirectconnectionexistsbetweenthepharynxandtheexcretorycanal,whichfunctionsinthereleaseofliquidurine. Maleshaveasingle-lobedgonad,avasdeferens,andatailspecializedformating,whichincorporatesspicules.Hermaphroditeshavetwoovaries,oviducts,andspermatheca,andasingleuterus. AnatomicaldiagramofamaleC.elegans C.elegansneuronscontaindendriteswhichextendfromthecelltoreceiveneurotransmitters,andaprocessthatextendstothenervering(the"brain")forasynapticconnectionbetweenneurons.[20]ThebiggestdifferenceisthatC.eleganshasmotorexcitatoryandinhibitoryneurons,knownascholinergicandgabaergicneurons,whichsimplyactasfurtherregulationforthetinycreature.Theyhavenoinfluenceonthenervoussystembesidesregulatingneuronimpulses.[21] Gutgranules[edit] NumerousgutgranulesarepresentintheintestineofC.elegans,thefunctionsofwhicharestillnotfullyknown,asaremanyotheraspectsofthisnematode,despitethemanyyearsthatithasbeenstudied.ThesegutgranulesarefoundinalloftheRhabditidaorders.Theyareverysimilartolysosomesinthattheyfeatureanacidicinteriorandthecapacityforendocytosis,buttheyareconsiderablylarger,reinforcingtheviewoftheirbeingstorageorganelles. Aremarkablefeatureofthegranulesisthatwhentheyareobservedunderultravioletlight,theyreactbyemittinganintensebluefluorescence.Anotherphenomenonseenistermed'deathfluorescence'.Asthewormsdie,adramaticburstofbluefluorescenceisemitted.Thisdeathfluorescencetypicallytakesplaceinananteriortoposteriorwavethatmovesalongtheintestine,andisseeninbothyoungandoldworms,whethersubjectedtolethalinjuryorpeacefullydyingofoldage. Manytheorieshavebeenpositedonthefunctionsofthegutgranules,withearlieronesbeingeliminatedbylaterfindings.Theyarethoughttostorezincasoneoftheirfunctions.Recentchemicalanalysishasidentifiedthebluefluorescentmaterialtheycontainasaglycosylatedformofanthranilicacid(AA).TheneedforthelargeamountsofAAthemanygutgranulescontainisquestioned.OnepossibilityisthattheAAisantibacterialandusedindefenseagainstinvadingpathogens.Anotherpossibilityisthatthegranulesprovidephotoprotection;theburstsofAAfluorescenceentailtheconversionofdamagingUVlighttorelativelyharmlessvisiblelight.Thisisseenapossiblelinktothemelanin–containingmelanosomes.[22] Alateral(left)sideanatomicaldiagramofanadult-stageC.eleganshermaphrodite Reproduction[edit] Thehermaphroditicwormisconsideredtobeaspecializedformofself-fertilefemale,asitssomaisfemale.Thehermaphroditicgermlineproducesmalegametesfirst,andlayseggsthroughitsuterusafterinternalfertilization.HermaphroditesproducealltheirspermintheL4stage(150spermcellspergonadalarm)andthenproduceonlyoocytes.Thehermaphroditicgonadactsasanovotestiswithspermcellsbeingstoredinthesameareaofthegonadastheoocytesuntilthefirstoocytepushesthespermintothespermatheca(achamberwhereintheoocytesbecomefertilizedbythesperm).[23] Themalecaninseminatethehermaphrodite,whichwillpreferentiallyusemalesperm(bothtypesofspermarestoredinthespermatheca). Onceherecognizesahermaphroditeworm,themalenematodebeginstracingthehermaphroditewithhistailuntilhereachesthevulvalregion.Themalethenprobestheregionwithhisspiculestolocatethevulva,insertsthem,andreleasessperm.[24]ThespermofC.elegansisamoeboid,lackingflagellaandacrosomes.[25]Whenself-inseminated,thewild-typewormlaysabout300eggs.Wheninseminatedbyamale,thenumberofprogenycanexceed1,000.Hermaphroditesdonottypicallymatewithotherhermaphrodites.At20 °C,thelaboratorystrainofC.elegans(N2)hasanaveragelifespanaround2–3weeksandagenerationtimeof3to4days. C.eleganshasfivepairsofautosomesandonepairofsexchromosomes.SexinC.elegansisbasedonanX0sex-determinationsystem.HermaphroditesofC.eleganshaveamatchedpairofsexchromosomes(XX);theraremaleshaveonlyonesexchromosome(X0). Sexdetermination[edit] C.elegansaremostlyhermaphroditicorganisms,producingbothspermsandoocytes.[26]Malesdooccurinthepopulationinarateofapproximately1in200hermaphrodites,butthetwosexesarehighlydifferentiated.[27][28]Malesdifferfromtheirhermaphroditiccounterpartsinthattheyaresmallerandcanbeidentifiedthroughtheshapeoftheirtail.[28]C.elegansreproducethroughaprocesscalledandrodioecy.Thismeansthattheycanreproduceintwoways:eitherthroughself-fertilizationinhermaphroditesorthroughhermaphroditesbreedingwithmales.Malesareproducedthroughnon-disjunctionoftheXchromosomesduringmeiosis.Thewormsthatreproducethroughself-fertilizationareatriskforhighlinkagedisequilibrium,whichleadstolowergeneticdiversityinpopulationsandanincreaseinaccumulationofdeleteriousalleles.[29]InC.elegans,somaticsexdeterminationisattributedtothetra-1gene.[30]Thetra-1isagenewithintheTRA-1transcriptionfactorsexdeterminationpathwaythatisregulatedpost-transcriptionallyandworksbypromotingfemaledevelopment.[30]Inhermaphrodites(XX),therearehighlevelsoftra-1activity,whichproducesthefemalereproductivesystemandinhibitsmaledevelopment.[26][30]Atacertaintimeintheirlifecycle,onedaybeforeadulthood,hermaphroditescanbeidentifiedthroughtheadditionofavulvaneartheirtail.Inmales(XO),therearelowlevelsoftra-1activity,resultinginamalereproductivesystem.[30]Recentresearchhasshownthattherearethreeothergenes,fem-1,fem-2,andfem-3,thatnegativelyregulatetheTRA-1pathwayandactasthefinaldeterminerofsexinC.elegans.[26] Evolution[edit] ThesexdeterminationsysteminC.elegansisatopicthathasbeenofinteresttoscientistsforyears.[27]Sincetheyareusedasamodelorganism,anyinformationdiscoveredaboutthewaytheirsexdeterminationsystemmighthaveevolvedcouldfurtherthesameevolutionarybiologyresearchinotherorganisms.Afteralmost30yearsofresearch,scientistshavebeguntoputtogetherthepiecesintheevolutionofsuchasystem.[27]Whattheyhavediscoveredisthatthereisacomplexpathwayinvolvedthathasseverallayersofregulation.[27]ThecloselyrelatedorganismCaenorhabditisbriggsaehasbeenstudiedextensivelyanditswholegenomesequencehashelpedputtogetherthemissingpiecesintheevolutionofC.eleganssexdetermination.[27]Ithasbeendiscoveredthattwogeneshaveassimilated,leadingtotheproteinsXOL-1andMIX-1havinganeffectonsexdeterminationinC.elegansaswell.[27]MutationsintheXOL-1pathwayleadstofeminizationinC.elegans.[31]Themix-1geneisknowntohypoactivatetheXchromosomeandregulatesthemorphologyofthemaletailinC.elegans.[32]Lookingatthenematodeasawhole,themaleandhermaphroditesexlikelyevolvedfromparallelevolution.[27]Parallelevolutionisdefinedassimilartraitsevolvingfromanancestorinsimilarconditions;simplyput,thetwospeciesevolveinsimilarwaysovertime.Anexampleofthiswouldbemarsupialandplacentalmammals.Scientistshavealsohypothesizedthathermaphroditeasexualreproduction,or"selfing",couldhaveevolvedconvergentlybystudyingspeciessimilartoC.elegans[27]Otherstudiesonthesexdeterminationevolutionsuggestthatgenesinvolvingspermevolveatthefasterratethanfemalegenes.[33]However,spermgenesontheXchromosomehavereducedevolutionrates.Spermgeneshaveshortcodingsequences,highcodonbias,anddisproportionaterepresentationamongorphangenes.[33]Thesecharacteristicsofspermgenesmaybethereasonfortheirhighratesofevolutionandmayalsosuggesthowspermgenesevolvedoutofhermaphroditeworms.Overall,scientistshaveageneralideaofthesexdeterminationpathwayinC.elegans,however,theevolutionofhowthispathwaycametobeisnotyetwelldefined. Development[edit] C.elegansembryonicdevelopment Embryonicdevelopment[edit] Thefertilizedzygoteundergoesrotationalholoblasticcleavage. Spermentryintotheoocytecommencesformationofananterior-posterioraxis.[34]Thespermmicrotubuleorganizingcenterdirectsthemovementofthespermpronucleustothefutureposteriorpoleoftheembryo,whilealsoincitingthemovementofPARproteins,agroupofcytoplasmicdeterminationfactors,totheirproperrespectivelocations.[35]AsaresultofthedifferenceinPARproteindistribution,thefirstcelldivisionishighlyasymmetric.[36]C.elegansembryogenesisisamongthebestunderstoodexamplesofasymmetriccelldivision.[37] Allcellsofthegermlinearisefromasingleprimordialgermcell,calledtheP4cell,establishedearlyinembryogenesis.[38][39]Thisprimordialcelldividestogeneratetwogermlineprecursorsthatdonotdividefurtheruntilafterhatching.[39] Axisformation[edit] TheresultingdaughtercellsofthefirstcelldivisionarecalledtheABcell(containingPAR-6andPAR-3)andtheP1cell(containingPAR-1andPAR-2).AsecondcelldivisionproducestheABpandABacellsfromtheABcell,andtheEMSandP2cellsfromtheP1cell.Thisdivisionestablishesthedorsal-ventralaxis,withtheABpcellformingthedorsalsideandtheEMScellmarkingtheventralside.[40]ThroughWntsignaling,theP2cellinstructstheEMScelltodividealongtheanterior-posterioraxis.[41]ThroughNotchsignaling,theP2celldifferentiallyspecifiestheABpandABacells,whichfurtherdefinesthedorsal-ventralaxis.Theleft-rightaxisalsobecomesapparentearlyinembryogenesis,althoughitisunclearexactlywhenspecificallytheaxisisdetermined.However,mosttheoriesoftheL-RaxisdevelopmentinvolvesomekindofdifferencesincellsderivedfromtheABcell.[42] Gastrulation[edit] Gastrulationoccursaftertheembryoreachesthe24-cellstage.[43]C.elegansareaspeciesofprotostomes,sotheblastoporeeventuallyformsthemouth.Involutionintotheblastoporebeginswithmovementoftheendodermcellsandsubsequentformationofthegut,followedbytheP4germlineprecursor,andfinallythemesodermcells,includingthecellsthateventuallyformthepharynx.Gastrulationendswhenepibolyofthehypoblastsclosestheblastopore.[44] Post-embryonicdevelopment[edit] Underenvironmentalconditionsfavourableforreproduction,hatchedlarvaedevelopthroughfourlarvalstages-L1,L2,L3,andL4-injust3daysat20 °C.Whenconditionsarestressed,asinfoodinsufficiency,excessivepopulationdensityorhightemperature,C.eleganscanenteranalternativethirdlarvalstage,L2d,calledthedauerstage(DauerisGermanforpermanent).Aspecificdauerpheromoneregulatesentryintothedauerstate.Thispheromoneiscomposedofsimilarderivativesofthe3,6-dideoxysugar,ascarylose.Ascarosides,namedaftertheascarylosebase,areinvolvedinmanysex-specificandsocialbehaviors.[45]Inthisway,theyconstituteachemicallanguagethatC.elegansusestomodulatevariousphenotypes.Dauerlarvaearestress-resistant;theyarethinandtheirmouthsaresealedwithacharacteristicdauercuticleandcannottakeinfood.Theycanremaininthisstageforafewmonths.[46][47]Thestageendswhenconditionsimprovefavourfurthergrowthofthelarva,nowmoultingintotheL4stage,eventhoughthegonaddevelopmentisarrestedattheL2stage.[48] Eachstagetransitionispunctuatedbyamoltoftheworm'stransparentcuticle.Transitionsthroughthesestagesarecontrolledbygenesoftheheterochronicpathway,anevolutionarilyconservedsetofregulatoryfactors.[49]ManyheterochronicgenescodeformicroRNAs,whichrepresstheexpressionofheterochronictranscriptionfactorsandotherheterochronicmiRNAs.[50]miRNAswereoriginallydiscoveredinC.elegans.[51]Importantdevelopmentaleventscontrolledbyheterochronicgenesincludethedivisionandeventualsyncitialfusionofthehypodermicseamcells,andtheirsubsequentsecretionofthealaeinyoungadults.Itisbelievedthattheheterochronicpathwayrepresentsanevolutionarilyconservedpredecessortocircadianclocks.[52] Somenematodeshaveafixed,geneticallydeterminednumberofcells,aphenomenonknownaseutely.TheadultC.eleganshermaphroditehas959somaticcellsandthemalehas1033cells,[53][54][55]althoughithasbeensuggestedthatthenumberoftheirintestinalcellscanincreasebyonetothreeinresponsetogutmicrobesexperiencedbymothers.[56]Muchoftheliteraturedescribesthecellnumberinmalesas1031,butthediscoveryofapairofleftandrightMCMneuronsincreasedthenumberbytwoin2015.[55]Thenumberofcellsdoesnotchangeaftercelldivisionceasesattheendofthelarvalperiod,andsubsequentgrowthisduesolelytoanincreaseinthesizeofindividualcells.[57] Ecology[edit] Mainarticle:HostmicrobeinteractionsinCaenorhabditiselegans ThedifferentCaenorhabditisspeciesoccupyvariousnutrient-andbacteria-richenvironments.Theyfeedonthebacteriathatdevelopindecayingorganicmatter(microbivory).Soillacksenoughorganicmattertosupportself-sustainingpopulations.C.eleganscansurviveonadietofavarietyofbacteria,butitswildecologyislargelyunknown.Mostlaboratorystrainsweretakenfromartificialenvironmentssuchasgardensandcompostpiles.Morerecently,C.eleganshasbeenfoundtothriveinotherkindsoforganicmatter,particularlyrottingfruit.[58] C.eleganscanalsousedifferentspeciesofyeast,includingCryptococcuslaurentiiandC.kuetzingii,assolesourcesoffood.[59]Althoughabacterivore,C.eleganscanbekilledbyanumberofpathogenicbacteria,includinghumanpathogenssuchasStaphylococcusaureus,[60]Pseudomonasaeruginosa,[61]SalmonellaentericaorEnterococcusfaecalis.[62] Invertebratessuchasmillipedes,insects,isopods,andgastropodscantransportdauerlarvaetovarioussuitablelocations.Thelarvaehavealsobeenseentofeedontheirhostswhentheydie.[63] Nematodescansurvivedesiccation,andinC.elegans,themechanismforthiscapabilityhasbeendemonstratedtobelateembryogenesisabundantproteins.[64] C.elegans,asothernematodes,canbeeatenbypredatornematodesandotheromnivores,includingsomeinsects.[65] TheOrsayvirusisavirusthataffectsC.elegans,aswellastheCaenorhabditiselegansCer1virus[66]andtheCaenorhabditiselegansCer13virus. Interactionswithfungi WildisolatesofCaenorhabditiselegansareregularlyfoundwithinfectionsbyMicrosporidiafungi.Onesuchspecies,Nematocidaparisii,replicatesintheintestinesofC.elegans.[67] Arthrobotrysoligosporaisthemodelorganismforinteractionsbetweenfungiandnematodes.[68]Itisthemostcommonandwidespreadnematodecapturingfungus. Useasamodelorganism[edit] Furtherinformation:HistoryofresearchonCaenorhabditiselegans Asymmetriccelldivisionsduringearlyembryogenesisofwild-typeC.elegans In1963,SydneyBrennerproposedusingC.elegansasamodelorganismfortheinvestigationprimarilyofneuraldevelopmentinanimals.Itisoneofthesimplestorganismswithanervoussystem.Theneuronsdonotfireactionpotentials,anddonotexpressanyvoltage-gatedsodiumchannels.[69]Inthehermaphrodite,thissystemcomprises302neurons[70]thepatternofwhichhasbeencomprehensivelymapped,[71]inwhatisknownasaconnectome,andshowntobeasmall-worldnetwork.[72] ResearchhasexploredtheneuralandmolecularmechanismsthatcontrolseveralbehaviorsofC.elegans,includingchemotaxis,thermotaxis,mechanotransduction,learning,memory,andmatingbehaviour.[73]In2019theconnectomeofthemalewaspublishedusingatechniquedistinctfromthatusedforthehermaphrodite.Thesamepaperusedthenewtechniquetoredothehermaphroditeconnectome,finding1,500newsynapses.[74] Ithasbeenusedasamodelorganismtostudymolecularmechanismsinmetabolicdiseases.[75] Brenneralsochoseitasitiseasytogrowinbulkpopulations,andconvenientforgeneticanalysis.[76]Itisamulticellulareukaryoticorganism,yetsimpleenoughtobestudiedingreatdetail.ThetransparencyofC.elegansfacilitatesthestudyofcellulardifferentiationandotherdevelopmentalprocessesintheintactorganism.Thespiculesinthemaleclearlydistinguishmalesfromfemales.Strainsarecheaptobreedandcanbefrozen.Whensubsequentlythawed,theyremainviable,allowinglong-termstorage.[13]Maintenanceiseasywhencomparedtoothermulticellularmodelorganisms.Afewhundrednematodescanbekeptonasingleagarplateandsuitablegrowthmedium.BrennerdescribedtheuseofamutantofE.coli–OP50.OP50isauracil-requiringorganismanditsdeficiencyintheplatepreventstheovergrowthofbacteriawhichwouldobscuretheworms.[77]TheuseofOP50doesnotdemandanymajorlaboratorysafetymeasures,sinceitisnon-pathogenicandeasilygrowninLuria-Bertani(LB)mediaovernight.[78] Celllineagemapping[edit] Thedevelopmentalfateofeverysinglesomaticcell(959intheadulthermaphrodite;1031intheadultmale)hasbeenmapped.[79][80]Thesepatternsofcelllineagearelargelyinvariantbetweenindividuals,whereasinmammals,celldevelopmentismoredependentoncellularcuesfromtheembryo. Asmentionedpreviously,thefirstcelldivisionsofearlyembryogenesisinC.elegansareamongthebestunderstoodexamplesofasymmetriccelldivisions,andthewormisaverypopularmodelsystemforstudyingdevelopmentalbiology.[37] Programmedcelldeath[edit] Programmedcelldeath(apoptosis)eliminatesmanyadditionalcells(131inthehermaphrodite,mostofwhichwouldotherwisebecomeneurons);this"apoptoticpredictability"hascontributedtotheelucidationofsomeapoptoticgenes.Celldeath-promotinggenesandasinglecell-deathinhibitorhavebeenidentified.[81] RNAinterferenceandgenesilencing[edit] Wild-typeC.eleganshermaphroditestainedwiththefluorescentdyeTexasRedtohighlightthenucleiofallcells RNAinterference(RNAi)isarelativelystraightforwardmethodofdisruptingthefunctionofspecificgenes.Silencingthefunctionofagenecansometimesallowaresearchertoinferitspossiblefunction.Thenematodecanbesoakedin,injectedwith,[82]orfedwithgeneticallytransformedbacteriathatexpressthedouble-strandedRNAofinterest,thesequenceofwhichcomplementsthesequenceofthegenethattheresearcherwishestodisable.[83] RNAihasemergedasapowerfultoolinthestudyoffunctionalgenomics.C.eleganshasbeenusedtoanalysegenefunctionsandclaimthepromiseoffuturefindingsinthesystematicgeneticinteractions.[84] EnvironmentalRNAiuptakeismuchworseinotherspeciesofwormsinthegenusCaenorhabditis.AlthoughinjectingRNAintothebodycavityoftheanimalinducesgenesilencinginmostspecies,onlyC.elegansandafewotherdistantlyrelatednematodescantakeupRNAfromthebacteriatheyeatforRNAi.[85]Thisabilityhasbeenmappeddowntoasinglegene,sid-2,which,wheninsertedasatransgeneinotherspecies,allowsthemtotakeupRNAforRNAiasC.elegansdoes.[86] Celldivisionandcellcycle[edit] Researchintomeiosishasbeenconsiderablysimplifiedsinceeverygermcellnucleusisatthesamegivenpositionasitmovesdownthegonad,soisatthesamestageinmeiosis.Inanearlyphaseofmeiosis,theoocytesbecomeextremelyresistanttoradiationandthisresistancedependsonexpressionofgenesrad51andatmthathavekeyrolesinrecombinationalrepair.[87][88]Genemre-11alsoplaysacrucialroleinrecombinationalrepairofDNAdamageduringmeiosis.[89]AstudyofthefrequencyofoutcrossinginnaturalpopulationsshowedthatselfingisthepredominantmodeofreproductioninC.elegans,butthatinfrequentoutcrossingeventsoccurataratearound1%.[90]Meiosesthatresultinselfingareunlikelytocontributesignificantlytobeneficialgeneticvariability,butthesemeiosesmayprovidetheadaptivebenefitofrecombinationalrepairofDNAdamagesthatarise,especiallyunderstressfulconditions.[91] Drugabuseandaddiction[edit] NicotinedependencecanalsobestudiedusingC.elegansbecauseitexhibitsbehavioralresponsestonicotinethatparallelthoseofmammals.Theseresponsesincludeacuteresponse,tolerance,withdrawal,andsensitization.[92] Biologicaldatabases[edit] Asformostmodelorganisms,scientiststhatworkinthefieldcurateadedicatedonlinedatabaseandtheWormBaseisthatforC.elegans.TheWormBaseattemptstocollateallpublishedinformationonC.elegansandotherrelatednematodes.Theirwebsitehasadvertisedarewardof$4000forthefinderofanewspeciesofcloselyrelatednematode.[93]Suchadiscoverywouldbroadenresearchopportunitieswiththeworm.[94] Ageing[edit] C.eleganshasbeenamodelorganismforresearchintoageing;forexample,theinhibitionofaninsulin-likegrowthfactorsignalingpathwayhasbeenshowntoincreaseadultlifespanthreefold;[95][96]whileglucosefeedingpromotesoxidativestressandreduceadultlifespanbyahalf.[75]Similarly,induceddegradationofaninsulin/IGF-1receptorlateinlifeextendedlifeexpectancyofwormsdramatically.[97] Long-livedmutantsofC.elegansweredemonstratedtoberesistanttooxidativestressandUVlight.[98]Theselong-livedmutantshadahigherDNArepaircapabilitythanwild-typeC.elegans.[98]KnockdownofthenucleotideexcisionrepairgeneXpa-1increasedsensitivitytoUVandreducedthelifespanofthelong-livedmutants.ThesefindingsindicatethatDNArepaircapabilityunderlieslongevity. ThecapacitytorepairDNAdamagebytheprocessofnucleotideexcisionrepairdeclineswithage.[99] C.elegansexposedto5mMlithiumchloride(LiCl)showedlengthenedlifespans.[100]Whenexposedto10μMLiCl,reducedmortalitywasobserved,butnotwith1μM.[101] C.eleganshasbeeninstrumentalintheidentificationofthefunctionsofgenesimplicatedinAlzheimer'sdisease,suchaspresenilin.[102]Moreover,extensiveresearchonC.eleganshasidentifiedRNA-bindingproteinsasessentialfactorsduringgermlineandearlyembryonicdevelopment.[103] Telomeres,thelengthofwhichhavebeenshowntocorrelatewithincreasedlifespananddelayedonsetofsenescenceinamultitudeoforganisms,fromC.elegans[104][105]tohumans,[106]showaninterestingbehaviourinC.elegans.WhileC.elegansmaintainsitstelomeresinacanonicalwaysimilartoothereukaryotes,incontrastDrosophilamelanogasterisnoteworthyinitsuseofretrotransposonstomaintainitstelomeres,[107]duringknock-outofthecatalyticsubunitofthetelomerase(trt-1)C.eleganscangaintheabilityofalternativetelomerelengthening(ALT).C.eleganswasthefirsteukaryotetogainALTfunctionalityafterknock-outofthecanonicaltelomerasepathway.[108]ALTisalsoobservedinabout10-15%ofallclinicalcancers.[109]ThusC.elegansisaprimecandidateforALTresearch.[110][111][112]Bayatetal.showedtheparadoxicalshorteningoftelomeresduringtrt-1over-expressionwhichleadtonearsterilitywhilethewormsevenexhibitedaslightincreaseinlifespan,despiteshortenedtelomeres.[113] Sleep[edit] C.elegansisnotableinanimalsleepstudiesasthemostprimitiveorganismtodisplaysleep-likestates.InC.elegans,alethargusphaseoccursshortlybeforeeachmoult.[114]C.eleganshasalsobeendemonstratedtosleepafterexposuretophysicalstress,includingheatshock,UVradiation,andbacterialtoxins.[115] Sensorybiology[edit] Whilethewormhasnoeyes,ithasbeenfoundtobesensitivetolightduetoathirdtypeoflight-sensitiveanimalphotoreceptorprotein,LITE-1,whichis10to100timesmoreefficientatabsorbinglightthantheothertwotypesofphotopigments(opsinsandcryptochromes)foundintheanimalkingdom.[116] C.elegansisremarkablyadeptattoleratingacceleration.Itcanwithstand400,000g's,accordingtogeneticistsattheUniversityofSãoPauloinBrazil.Inanexperiment,96%ofthemwerestillalivewithoutadverseeffectsafteranhourinanultracentrifuge.[117] Spaceflightresearch[edit] C.elegansmadenewswhenspecimenswerediscoveredtohavesurvivedtheSpaceShuttleColumbiadisasterinFebruary2003.[118]Later,inJanuary2009,livesamplesofC.elegansfromtheUniversityofNottinghamwereannouncedtobespendingtwoweeksontheInternationalSpaceStationthatOctober,inaspaceresearchprojecttoexploretheeffectsofzerogravityonmuscledevelopmentandphysiology.Theresearchwasprimarilyaboutgeneticbasisofmuscleatrophy,whichrelatestospaceflightorbeingbed-ridden,geriatric,ordiabetic.[119]DescendantsofthewormsaboardColumbiain2003werelaunchedintospaceonEndeavourfortheSTS-134mission.[120]AdditionalexperimentsonmuscledystrophyduringspaceflightwillbecarriedonboardoftheISSstartinginDecember2018.[121][needsupdate] Genetics[edit] Genome[edit] GenomicinformationKaryotypeofC.elegansexplanationofcolors MitoticchromosomesofC.elegans.DNA(red)/Kinetochores(green).Holocentricorganisms,includingC.elegans,assemblediffusekinetochoresalongtheentirepolewardfaceofeachsisterchromatid.NCBIgenomeID41PloidydiploidGenomesize101.169Mb(haploid)Numberofchromosomes5pairsofautosomes(I,II,III,IVandV)+1or2sexchromosomes(X[122])Yearofcompletion1998SequencedorganellemitochondrionOrganellesize0,01Mb C.eleganshermaphrodite C.eleganswasthefirstmulticellularorganismtohaveitswholegenomesequenced.Thesequencewaspublishedin1998,[123]althoughsomesmallgapswerepresent;thelastgapwasfinishedbyOctober2002.[citationneeded]IntherunuptothewholegenometheC.elegansSequencingConsortium/C.elegansGenomeProjectreleasedseveralpartialscansincludingWilsonetal.1994.[124][125][126] Sizeandgenecontent[edit] TheC.elegansgenomeisabout100millionbasepairslongandconsistsofsixpairsofchromosomesinhermaphroditesorfivepairsofautosomeswithXOchromosomeinmaleC.elegansandamitochondrialgenome.Itsgenedensityisaboutonegeneperfivekilo-basepairs.Intronsmakeup26%andintergenicregions47%ofthegenome.Manygenesarearrangedinclustersandhowmanyoftheseareoperonsisunclear.[127]C.elegansandothernematodesareamongthefeweukaryotescurrentlyknowntohaveoperons;theseincludetrypanosomes,flatworms(notablythetrematodeSchistosomamansoni),andaprimitivechordatetunicateOikopleuradioica.Manymoreorganismsarelikelytobeshowntohavetheseoperons.[128] Thegenomecontainsanestimated20,470protein-codinggenes.[129]About35%ofC.elegansgeneshavehumanhomologs.Remarkably,humangeneshavebeenshownrepeatedlytoreplacetheirC.eleganshomologswhenintroducedintoC.elegans.Conversely,manyC.elegansgenescanfunctionsimilarlytomammaliangenes.[46] ThenumberofknownRNAgenesinthegenomehasincreasedgreatlyduetothe2006discoveryofanewclasscalled21U-RNAgenes,[130]andthegenomeisnowbelievedtocontainmorethan16,000RNAgenes,upfromasfewas1,300in2005.[131] Scientificcuratorscontinuetoappraisethesetofknowngenes;newgenemodelscontinuetobeaddedandincorrectonesmodifiedorremoved. ThereferenceC.elegansgenomesequencecontinuestochangeasnewevidencerevealserrorsintheoriginalsequencing.Mostchangesareminor,addingorremovingonlyafewbasepairsofDNA.Forexample,theWS202releaseofWormBase(April2009)addedtwobasepairstothegenomesequence.[132]Sometimes,moreextensivechangesaremadeasnotedintheWS197releaseofDecember2008,whichaddedaregionofover4,300bptothesequence.[133][134] TheC.elegansGenomeProject'sWilsonetal.1994foundCelVav[126]andavonWillebrandfactorAdomain[124]andwithWilsonetal.1998providesthefirstcredibleevidenceforanarylhydrocarbonreceptor(AHR)homologoutsideofvertebrates.[125]2 Relatedgenomes[edit] In2003,thegenomesequenceoftherelatednematodeC.briggsaewasalsodetermined,allowingresearcherstostudythecomparativegenomicsofthesetwoorganisms.[135]Thegenomesequencesofmorenematodesfromthesamegenuse.g.,C.remanei,[136]C.japonica[137]andC.brenneri(namedafterBrenner),havealsobeenstudiedusingtheshotgunsequencingtechnique.[138]Thesesequenceshavenowbeencompleted.[139][140] Othergeneticstudies[edit] C.elegansadultwithGFPcodingsequenceinsertedintoahistone-encodinggenebyCas9-triggeredhomologousrecombination Asof2014,C.elegansisthemostbasalspeciesinthe'Elegans'group(10species)ofthe'Elegans'supergroup(17species)inphylogeneticstudies.Itformsabranchofitsowndistincttoanyotherspeciesofthegroup.[141] Tc1transposonisaDNAtransposonactiveinC.elegans. Scientificcommunity[edit] In2002,theNobelPrizeinPhysiologyorMedicinewasawardedtoSydneyBrenner,H.RobertHorvitz,andJohnSulstonfortheirworkonthegeneticsoforgandevelopmentandprogrammedcelldeathinC.elegans.The2006NobelPrizeinPhysiologyorMedicinewasawardedtoAndrewFireandCraigC.MellofortheirdiscoveryofRNAinterferenceinC.elegans.[142]In2008,MartinChalfiesharedaNobelPrizeinChemistryforhisworkongreenfluorescentprotein;someoftheresearchinvolvedtheuseofC.elegans. ManyscientistswhoresearchC.eleganscloselyconnecttoSydneyBrenner,withwhomalmostallresearchinthisfieldbeganinthe1970s;theyhaveworkedaseitherapostdoctoralorapostgraduateresearcherinBrenner'slaborinthelabofsomeonewhopreviouslyworkedwithBrenner.Mostwhoworkedinhislablaterestablishedtheirownwormresearchlabs,therebycreatingafairlywell-documented"lineage"ofC.elegansscientists,whichwasrecordedintotheWormBasedatabaseinsomedetailatthe2003InternationalWormMeeting.[143] Seealso[edit] WikimediaCommonshasmediarelatedtoCaenorhabditiselegans. 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Furtherreading[edit] BirdJ,BirdAC(1991).Thestructureofnematodes.AcademicPress.pp. 1,69–70,152–153,165,224–225.ISBN 978-0-12-099651-3. Hope,IA(1999).C.elegans:apracticalapproach.OxfordUniversityPress.pp. 1–6.ISBN 978-0-19-963738-6. RiddleDL,BlumenthalT,MeyerRJ,PriessJR(1997).C.elegansII.ColdSpringHarborLaboratoryPress.pp. 1–4,679–683.ISBN 978-0-87969-532-3. Externallinks[edit] WikimediaCommonshasmediarelatedto: Caenorhabditiselegans(category) WikispecieshasinformationrelatedtoCaenorhabditiselegans. ScholiahasataxonprofileforCaenorhabditiselegans. ScholiahasatopicprofileforCaenorhabditiselegans. BrennerS(2002)Nature'sGifttoScience.In.http://nobelprize.org/nobel_prizes/medicine/laureates/2002/brenner-lecture.pdf(alsoHorvitzandSulstonlectures) WormBase–anextensiveonlinedatabasecoveringthebiologyandgenomicsofC.elegansandothernematodes WormAtlas–onlinedatabaseonallaspectsofC.elegansanatomywithdetailedexplanationsandhigh-qualityimages WormBook–onlinereviewofC.elegansbiology AceViewWormGenes–anothergenomedatabaseforC.elegans,maintainedattheNCBI C.elegansII–afreeonlinetextbook. WormWebNeuralNetwork–anonlinetoolforvisualizingandnavigatingtheconnectomeofC.elegans C.elegansmovies–avisualintroductiontoC.elegans Viewthece11genomeassemblyintheUCSCGenomeBrowser. Caenorhabditiselegansateppo.int(EPPOcodeCAEOEL) vteMajormodelorganismsingenetics Lambdaphage E.coli Chlamydomonas Tetrahymena Buddingyeast Fissionyeast Neurospora Maize Arabidopsis Medicagotruncatula C.elegans Drosophila Xenopus Zebrafish Rat Mouse Taxonidentifiers Wikidata:Q91703 Wikispecies:Caenorhabditiselegans BioLib:88399 BOLD:27260 CoL:87THG EoL:403869 EPPO:CAEOEL EUNIS:247866 FaunaEuropaea:224245 FaunaEuropaea(new):6dd1ded7-802e-4779-877e-e20eafcada89 GBIF:2283683 iNaturalist:372486 IRMNG:10955681 ITIS:63332 NBN:NHMSYS0000068531 NCBI:6239 SeaLifeBase:35925 WoRMS:583290 Retrievedfrom"https://en.wikipedia.org/w/index.php?title=Caenorhabditis_elegans&oldid=1103916091" Categories:CaenorhabditiselegansNematodesdescribedin1900AnimalmodelsinneuroscienceHiddencategories:WebarchivetemplatewaybacklinksArticleswithshortdescriptionShortdescriptionmatchesWikidataGoodarticlesArticleswith'species'microformatsArticlescontainingpotentiallydatedstatementsfrom2019AllarticlescontainingpotentiallydatedstatementsWikipediaarticlesinneedofupdatingfromJanuary2021AllWikipediaarticlesinneedofupdatingAllarticleswithunsourcedstatementsArticleswithunsourcedstatementsfromJanuary2022CommonscategorylinkfromWikidataArticlescontainingvideoclips Navigationmenu Personaltools NotloggedinTalkContributionsCreateaccountLogin Namespaces ArticleTalk English Views ReadEditViewhistory More Search Navigation MainpageContentsCurrenteventsRandomarticleAboutWikipediaContactusDonate Contribute HelpLearntoeditCommunityportalRecentchangesUploadfile Tools WhatlinkshereRelatedchangesUploadfileSpecialpagesPermanentlinkPageinformationCitethispageWikidataitem Print/export DownloadasPDFPrintableversion Inotherprojects WikimediaCommonsWikispecies Languages العربيةBosanskiCatalàCebuanoČeštinaDanskDeutschEestiEspañolEsperantoEuskaraفارسیFrançaisGalego客家語/Hak-kâ-ngî한국어BahasaIndonesiaItalianoעבריתLatinaMagyarمصرىNederlands日本語NorskbokmålNorsknynorskPolskiPortuguêsРусскийScotsSicilianuSimpleEnglishSlovenščinaСрпски/srpskiSrpskohrvatski/српскохрватскиSuomiSvenskaTagalogไทยTürkçeУкраїнськаئۇيغۇرچە/UyghurcheTiếngViệtWinaray粵語中文 Editlinks