Adenoviridae - Wikipedia

Adenoviruses are medium-sized (90–100 nm), nonenveloped viruses with an icosahedral nucleocapsid containing a double-stranded DNA genome. Adenoviridae FromWikipedia,thefreeencyclopedia Jumptonavigation Jumptosearch Familyofviruses "ad5","ad26","ad35",and"ad48"redirecthere.Fortheyears,seeAD5,AD26,AD35,andAD48.Forotheruses,seeAD-5(disambiguation),AD26(disambiguation),andAD35(disambiguation). Adenoviruses Transmissionelectronmicrographoftwoadenovirusparticles Virusclassification (unranked): Virus Realm: Varidnaviria Kingdom: Bamfordvirae Phylum: Preplasmiviricota Class: Tectiliviricetes Order: Rowavirales Family: Adenoviridae Genera Atadenovirus Aviadenovirus Ichtadenovirus Mastadenovirus Siadenovirus Testadenovirus AdenovirusD26structuralmodelatatomicresolution[1] Adenoviruses(membersofthefamilyAdenoviridae)aremedium-sized(90–100nm),nonenveloped(withoutanouterlipidbilayer)viruseswithanicosahedralnucleocapsidcontainingadouble-strandedDNAgenome.[2]Theirnamederivesfromtheirinitialisolationfromhumanadenoidsin1953.[3] Theyhaveabroadrangeofvertebratehosts;inhumans,morethan50distinctadenoviralserotypeshavebeenfoundtocauseawiderangeofillnesses,frommildrespiratoryinfectionsinyoungchildren(knownasthecommoncold)tolife-threateningmulti-organdiseaseinpeoplewithaweakenedimmunesystem.[2] Contents 1Virology 1.1Classification 1.2Diversity 1.3Structure 1.4Genome 1.5Replication 1.6Multiplicityreactivation 2Epidemiology 2.1Transmission 2.2Humans 2.3Animals 3Prevention 4Diagnosis 5Infections 6Treatment 7Useingenetherapyandvaccination 7.1Genetherapy 7.2Vaccines 7.2.1HIVinfectionconcerns 8Seealso 9References 10Externallinks Virology[edit] Classification[edit] Thisfamilycontainsthefollowinggenera:[4] Atadenovirus Aviadenovirus Ichtadenovirus Mastadenovirus(includingallhumanadenoviruses) Siadenovirus Testadenovirus Diversity[edit] Inhumans,currentlythereare88humanadenoviruses(HAdVs)insevenspecies(HumanadenovirusAtoG):[5] A:12,18,31 B:3,7,11,14,16,21,34,35,50,55 C:1,2,5,6,57[6] D:8,9,10,13,15,17,19,20,22,23,24,25,26,27,28,29,30,32,33,36,37,38,39,42,43,44,45,46,47,48,49,51,53,54,56,[7]58,59,60,62,63,[8]64,65,67,69,[9]70,71,72,73,74,75 E:4 F:40,41[10] G:52[11] Differenttypes/serotypesareassociatedwithdifferentconditions:[12] respiratorydisease(mainlyspeciesHAdV-BandC) conjunctivitis(HAdV-BandD) gastroenteritis(HAdV-Ftypes40,41,HAdV-Gtype52) obesityoradipogenesis(HAdV-Atype31,HAdV-Ctype5,HAdV-Dtypes9,36,37)[13] AllthesetypesarecalledHumanmastadenovirusA–GbytheICTV,becauseallaremembersofthegenusMastadenovirus. Structure[edit] Thestructureofadenovirus.1=pentoncapsomers,2=hexoncapsomers,and3=viralgenome(lineardsDNA) Adenovirusesaremedium-sized(90–100 nm).[2]Thevirionsarecomposedofonelinearpieceofdouble-strandedDNAinsideanicosahedralcapsid.240hexonproteinsmakeupthebulkofthecapsid,whiletwelvepentonbasescaptheicosahedron'scorners.Thepentonbasesareassociatedwithprotrudingfibersthataidinattachmenttothehostcellviathereceptoronitssurface.[14] In2010,thestructureofthehumanadenoviruswassolvedattheatomiclevel,makingitthelargesthigh-resolutionmodelever.Thevirusiscomposedofaround1millionaminoacidresiduesandweighsaround150MDa.[15][16] Genome[edit] Mainarticle:Adenovirusgenome Schematicdiagramofthelinearadenovirusgenome,showingEarlygenes(E)andLategenes(L). Theadenovirusgenomeislinear,non-segmenteddouble-stranded(ds)DNAthatisbetween26and48Kbp.[2]Thisallowsthevirustotheoreticallycarry22to40genes.AlthoughthisissignificantlylargerthanothervirusesinitsBaltimoregroup,itisstillaverysimplevirusandisheavilyreliantonthehostcellforsurvivalandreplication.Aninterestingfeatureofthisviralgenomeisthatithasaterminal55kDaproteinassociatedwitheachofthe5'endsofthelineardsDNA.Theseareusedasprimersinviralreplicationandensurethattheendsofthevirus'lineargenomeareadequatelyreplicated. Replication[edit] AdenovirusespossessalineardsDNAgenomeandareabletoreplicateinthenucleusofvertebratecellsusingthehost'sreplicationmachinery.[2]Entryofadenovirusesintothehostcellinvolvestwosetsofinteractionsbetweenthevirusandthehostcell.[2]Mostoftheactionoccursatthevertices.Entryintothehostcellisinitiatedbytheknobdomainofthefiberproteinbindingtothecellreceptor.[2]Thetwocurrentlyestablishedreceptorsare:CD46forthegroupBhumanadenovirusserotypesandthecoxsackievirus/adenovirusreceptor(CAR)forallotherserotypes.[2]TherearesomereportssuggestingMHCmoleculesandsialicacidresiduesfunctioninginthiscapacityaswell.Thisisfollowedbyasecondaryinteraction,whereamotifinthepentonbaseprotein(seecapsomere)interactswithanintegrinmolecule.Itistheco-receptorinteractionthatstimulatesentryoftheadenovirus.Thisco-receptormoleculeisαVintegrin.Bindingtoαvintegrinresultsinendocytosisofthevirusparticleviaclathrin-coatedpits.AttachmenttoαVintegrinstimulatescellsignalingandthusinducesactinpolymerization,whichfacilitatesclathrin-mediatedendocytosis,andresultsinvirion'sentryintothehostcellwithinanendosome.[17] Oncethevirushassuccessfullygainedentryintothehostcell,theendosomeacidifies,whichaltersvirustopologybycausingcapsidcomponentstodisband.ThecapsidisdestabilizedandproteinVI,whichisoneofthecapsidconstituents(seeAdenovirusgenome)isreleasedfromit.[18]Thesechanges,aswellasthetoxicnatureofthepentons,destroytheendosome,resultinginthemovementofthevirionintothecytoplasm.[2]Withthehelpofcellularmicrotubules,thevirusistransportedtothenuclearporecomplex,wherebytheadenovirusparticledisassembles.ViralDNAissubsequentlyreleased,whichcanenterthenucleusviathenuclearpore.[19]AfterthistheDNAassociateswithhistonemoleculesalreadypresentinthenucleus,whichallowsittointeractwiththehostcelltranscriptionmachinery[citationneeded].Then,viralgeneexpressioncanoccur,withoutintegratingtheviralgenomeintohostcellchromosomes,[20]andnewvirusparticlescanbegenerated. TheadenoviruslifecycleisseparatedbytheDNAreplicationprocessintotwophases:anearlyandalatephase.[2]Inbothphases,aprimarytranscriptthatisalternativelysplicedtogeneratemonocistronicmRNAscompatiblewiththehost'sribosomeisgenerated,allowingfortheproductstobetranslated. Theearlygenesareresponsibleforexpressingmainlynon-structural,regulatoryproteins.[2]Thegoaloftheseproteinsisthreefold:toaltertheexpressionofhostproteinsthatarenecessaryforDNAsynthesis;toactivateothervirusgenes(suchasthevirus-encodedDNApolymerase);andtoavoidprematuredeathoftheinfectedcellbythehost-immunedefenses(blockageofapoptosis,blockageofinterferonactivity,andblockageofMHCclassItranslocationandexpression). Someadenovirusesunderspecializedconditionscantransformcellsusingtheirearlygeneproducts.E1A(bindsRetinoblastomatumorsuppressorprotein)hasbeenfoundtoimmortalizeprimarycellsinvitroallowingE1B(bindsp53tumorsuppressor)toassistandstablytransformthecells.Nevertheless,theyarereliantuponeachothertosuccessfullytransformthehostcellandformtumors.E1AismostlyintrinsicallydisorderedproteinandcontainsCR3domainwhichiscriticalfortranscriptionalactivation.[21] DNAreplicationseparatestheearlyandlatephases.Oncetheearlygeneshaveliberatedadequatevirusproteins,replicationmachinery,andreplicationsubstrates,replicationoftheadenovirusgenomecanoccur.Aterminalproteinthatiscovalentlyboundtothe5'endoftheadenovirusgenomeactsasaprimerforreplication.TheviralDNApolymerasethenusesastranddisplacementmechanism,asopposedtotheconventionalOkazakifragmentsusedinmammalianDNAreplication,toreplicatethegenome. ThelatephaseoftheadenoviruslifecycleisfocusedonproducingsufficientquantitiesofstructuralproteintopackallthegeneticmaterialproducedbyDNAreplication.[2]Oncetheviralcomponentshavesuccessfullybeenreplicated,thevirusisassembledintoitsproteinshellsandreleasedfromthecellasaresultofvirallyinducedcelllysis.[2] Multiplicityreactivation[edit] Adenovirusiscapableofmultiplicityreactivation(MR)[22](YamamotoandShimojo,1971).MRistheprocessbywhichtwo,ormore,virusgenomescontaininglethaldamageinteractwithintheinfectedcelltoformaviablevirusgenome.SuchMRwasdemonstratedforadenovirus12aftervirionswereirradiatedwithUVlightandallowedtoundergomultipleinfectionofhostcells.[22]Inareview,numerousexamplesofMRindifferentvirusesweredescribed,anditwassuggestedthatMRisacommonformofsexualinteractionthatprovidesthesurvivaladvantageofrecombinationalrepairofgenomedamages.[23] Epidemiology[edit] Transmission[edit] AdenovirusesareunusuallystabletochemicalorphysicalagentsandadversepHconditions,allowingforprolongedsurvivaloutsideofthebodyandwater.Adenovirusesarespreadprimarilyviarespiratorydroplets,howevertheycanalsobespreadbyfecalroutes.Researchintothemolecularmechanismsunderlyingadenoviraltransmissionprovideempiricalevidenceinsupportofthehypothesisthatcoxsackievirus/adenovirusreceptors(CARs)areneededtotransportadenovirusesintocertainnaive/progenitorcelltypes.[24] Humans[edit] Mainarticle:Adenovirusinfection Humansinfectedwithadenovirusesdisplayawiderangeofresponses,fromnosymptomsatalltothesevereinfectionstypicalofAdenovirusserotype14. Animals[edit] Seealso:DA2PPCVaccine BatadenovirusTJM(Bt-AdV-TJM)isanovelspeciesoftheMastadenovirusgenusisolatedfromMyotisandScotophiluskuhliiinChina.[25]ItismostcloselyrelatedtothetreeshrewandcanineAdVs.[26] Twotypesofcanineadenovirusesarewellknown,type1and2.Type1(CAdV-1)causesinfectiouscaninehepatitis,apotentiallyfataldiseaseinvolvingvasculitisandhepatitis.Type1infectioncanalsocauserespiratoryandeyeinfections.CAdV-1alsoaffectsfoxes(VulpesvulpesandVulpeslagopus)andmaycausehepatitisandencephalitis.Canineadenovirus2(CAdV-2)isoneofthepotentialcausesofkennelcough.CorevaccinesfordogsincludeattenuatedliveCAdV-2,whichproducesimmunitytoCAdV-1andCAdV-2.CAdV-1wasinitiallyusedinavaccinefordogs,butcornealedemawasacommoncomplication.[27] Squirreladenovirus(SqAdV)isreportedtocauseenteritisinredsquirrelsinEurope,whilegraysquirrelsseemtoberesistant.SqAdVismostcloselyrelatedtotheadenovirusofguineapigs(GpAdV). Adenovirusinreptilesispoorlyunderstood,butresearchiscurrentlyinprogress. Adenovirusesarealsoknowntocauserespiratoryinfectionsinhorses,cattle,pigs,sheep,andgoats.Equineadenovirus1canalsocausefataldiseaseinimmunocompromisedArabianfoals,involvingpneumoniaanddestructionofpancreaticandsalivaryglandtissue.[27]Tupaiaadenovirus(TAV)(treeshrewadenovirus1)hasbeenisolatedfromtreeshrews. Otarineadenovirus1hasbeenisolatedfromsealions(Zalophuscalifornianus).[28] Thefowladenovirusesareassociatedwithmanydiseaseconditionsindomesticfowllikeinclusionbodyhepatitis,hydropericardiumsyndrome,[29]Eggdropsyndrome,Quailbronchitis,Gizzarderosionsandmanyrespiratoryconditions.Theyhavealsobeenisolatedfromwildblackkites(Milvusmigrans).[30] Titimonkeyadenoviruswasisolatedfromacolonyofmonkeys.[31] Prevention[edit] Mainarticle:Adenovirusvaccine Currentlythereisavaccineforadenovirustype4and7forUSmilitarypersonnelonly.USmilitarypersonnelaretherecipientsofthisvaccinebecausetheymaybeatahigherriskofinfection.[citationneeded]Thevaccinecontainsalivevirus,whichmaybeshedinstoolandleadtotransmission.Thevaccineisnotapprovedforuseoutsideofthemilitary,asithasnotbeentestedinstudiedinthegeneralpopulationoronpeoplewithweakenedimmunesystems.[32] Inthepast,USmilitaryrecruitswerevaccinatedagainsttwoserotypesofadenovirus,withacorrespondingdecreaseinillnessescausedbythoseserotypes.Thatvaccineisnolongermanufactured.TheU.S.ArmyMedicalResearchandMaterielCommandannouncedon31October2011thatanewadenovirusvaccine,whichreplacestheolderversionthathasbeenoutofproductionforoveradecade,wasshippedtobasictrainingsiteson18October2011.Moreinformationisavailablehere.[33] Preventionofadenovirus,aswellasotherrespiratoryillnesses,involvesfrequenthandwashingformorethan20seconds,avoidingtouchingtheeyes,face,andnosewithunwashedhands,andavoidingclosecontactwithpeoplewithsymptomaticadenovirusinfection.Thosewithsymptomaticadenovirusinfectionareadditionallyadvisedtocoughorsneezeintothearmorelbowinsteadofthehand,toavoidsharingcupsandeatingutensils,andtorefrainfromkissingothers.Chlorinationofswimmingpoolscanpreventoutbreaksofconjunctivitiscausedbyadenovirus.[32] Diagnosis[edit] Diagnosisisfromsymptomsandhistory.Testsareonlynecessaryinveryseriouscases.Testsincludebloodtests,eyes,noseorthroatswabs,stoolsampletests,andchestx-rays.[34]Inthelaboratory,adenoviruscanbeidentifiedwithantigendetection,polymerasechainreaction(PCR),virusisolationandserology.Evenifadenovirusisfoundtobepresent,itmaynotbethecauseofanysymptoms.Someimmunocompromisedindividualscanshedthevirusforweeksandshownosymptoms.[35] Infections[edit] Mainarticle:Adenovirusinfection Mostinfectionswithadenovirusresultininfectionsoftheupperrespiratorytract.Adenovirusinfectionsoftenpresentasconjunctivitis,tonsillitis(whichmaylookexactlylikestrepthroatandcannotbedistinguishedfromstrepexceptbythroatculture),anearinfection,orcroup.[36]Adenovirusestypes40and41canalsocausegastroenteritis.[37]Acombinationofconjunctivitisandtonsillitisisparticularlycommonwithadenovirusinfections. Somechildren(especiallytheyoungest)candevelopadenovirusbronchiolitisorpneumonia,bothofwhichcanbesevere.Inbabies,adenovirusescanalsocausecoughingfitsthatlookalmostexactlylikewhoopingcough.Adenovirusescanalsocauseviralmeningitisorencephalitis.Rarely,adenoviruscancausehemorrhagiccystitis(inflammationoftheurinarybladder—aformofurinarytractinfection—withbloodintheurine). Mostpeoplerecoverfromadenovirusinfectionsbythemselves,butpeoplewithimmunodeficiencysometimesdieofadenovirusinfections,and—rarely—evenpreviouslyhealthypeoplecandieoftheseinfections.[38]Thismaybebecausesometimesadenoviralinfectioncanleadtocardiacdisorders.Forexample,inonestudy,somecardiacsamplesofpatientswithdilatedcardiomyopathywerepositiveforpresenceofadenovirustype8.[39] Adenovirusesareoftentransmittedbyexpectoration(e.g.aerosols),buttheycanalsobetransmittedbycontactwithaninfectedperson,orbyvirusparticlesleftonobjectssuchastowelsandfaucethandles.Somepeoplewithadenovirusgastroenteritismayshedthevirusintheirstoolsformonthsaftergettingoverthesymptoms.Theviruscanbepassedthroughwaterinswimmingpoolsthatarenotsufficientlychlorinated. Aswithmanyotherillnesses,goodhandwashingpracticeisonewaytoinhibittheperson-to-persontransmissionofadenoviruses.Heatandbleachwillkilladenovirusesonobjects.[citationneeded] Treatment[edit] Therearenoprovenantiviraldrugstotreatadenoviralinfections,sotreatmentislargelydirectedatthesymptoms(suchasacetaminophenforfever).Theantiviraldrugcidofovirhashelpedcertainofthosepatientswhohadseverecasesofillness;thenumberhelpedandtowhatdegree,andtheparticularcomplicationsorsymptomsithelpedwith,andwhenandwherethishappened,werenotgiveninthesource.[40]Adoctormaygiveantibioticeyedropsforconjunctivitis,whileawaitingresultsofbacterialcultures,andtohelppreventsecondarybacterialinfections.Currently,thereisnoadenovirusvaccineavailabletothegeneralpublic,butavaccineisavailablefortheUnitedStatesmilitaryforTypes4and7. Useingenetherapyandvaccination[edit] Genetherapy[edit] Adenoviruseshavelongbeenapopularviralvectorforgenetherapyduetotheirabilitytoaffectbothreplicatingandnon-replicatingcells,accommodatelargetransgenes,andcodeforproteinswithoutintegratingintothehostcellgenome.[20]Morespecifically,theyareusedasavehicletoadministertargetedtherapy,[41]intheformofrecombinantDNAorprotein.Thistherapyhasbeenfoundespeciallyusefulintreatingmonogenicdisease(e.g.cysticfibrosis,X-linkedSCID,alpha1-antitrypsindeficiency)andcancer.[20]InChina,oncolyticadenovirusisanapprovedcancertreatment.[42]SpecificmodificationsonfiberproteinsareusedtotargetAdenovirustocertaincelltypes;[43]amajoreffortismadetolimithepatotoxicityandpreventmultipleorganfailure.Adenovirusdodecahedroncanqualifyasapotentdeliveryplatformforforeignantigenstohumanmyeloiddendriticcells(MDC),andthatitisefficientlypresentedbyMDCtoM1-specificCD8+Tlymphocytes.[44] AdenovirushasbeenusedfordeliveryofCRISPR/Cas9geneeditingsystems,buthighimmunereactivitytoviralinfectionhasposedchallengesinuseforpatients. Vaccines[edit] Furtherinformation:Viralvectorvaccine Modified(recombinant)adenovirusvectors,includingreplicationincompetenttypes,candeliverDNAcodingforspecificantigens.[45] AdenovirushavebeenusedtoproduceviralvectorCOVID-19vaccines."InfourcandidateCOVID-19vaccines...Ad5...servesasthe'vector'totransportthesurfaceproteingeneofSARS-CoV-2".[46]Thegoalistogeneticallyexpressthespikeglycoproteinofsevereacuterespiratorysyndromecoronavirus2(SARS-CoV-2).Areplication-deficientchimpanzeeadenovirusvaccinevector(ChAdOx1)isusedbytheOxford–AstraZenecaCOVID-19vaccinethathasbeenapprovedforuse.[47][48]TheJanssenCOVID-19vaccineusesmodifiedrecombinantadenovirustype-26(Ad26).[49]Recombinantadenovirustype-5(Ad5)arebeingusedbyAd5-nCoV,[50]ImmunityBioandUQ-CSLV451.TheGam-COVID-Vac(akaSputnik-V)productisinnovativebecauseanAd26basedvaccineisusedonthefirstdayandanAd5vaccineisusedonday21.[49]AnotheroneisChAd-SARS-CoV-2-S;thevaccinereportedlypreventedmicethatweregeneticallymodifiedtohavehumanACE2(hACE2)receptors,presumablyreceptorsthatallowvirus-entryintothecells,frombeinginfectedwithSARS-CoV-2.[51][52] PossibleissueswithusingAdenovirusasvaccinevectorsinclude:thehumanbodydevelopsimmunitytothevectoritself,makingsubsequentboostershotsdifficultorimpossible.[53]Insomecases,peoplehavepre-existingimmunitytoAdenoviruses,makingvectordeliveryineffective.[54] HIVinfectionconcerns[edit] Thissectionneedsmoremedicalreferencesforverificationorreliestooheavilyonprimarysources.Pleasereviewthecontentsofthesectionandaddtheappropriatereferencesifyoucan.Unsourcedorpoorlysourcedmaterialmaybechallengedandremoved.Findsources: "Adenoviridae" – news ·newspapers ·books ·scholar ·JSTOR(April2021) TheuseofAd5vaccinesforCOVID-19worriedresearcherswhohadexperiencewithtwofailedtrialsofanAd5vaccine,PhambiliandSTEP,duetotheincreasedriskforuncircumcisedmalepatientsofcontractingHIV-1viaunprotectedanalsex.[55]Atthetime,itwasconcludedthatheightenedriskofHIVreceptionmaybeobservedforanyAd5-basedvectorvaccine.[56]InOctober2020,theseresearcherswroteinTheLancet:"Onthebasisofthesefindings,weareconcernedthatuseofanAd5vectorforimmunisationagainstSARS-CoV-2couldsimilarlyincreasetheriskofHIV-1acquisitionamongmenwhoreceivethevaccine."[57][58]Vaccinesusingothertechnologieswouldnotbeaffected,butSputnikV,ConvideciaandImmunityBio'shAd5would.[59]TwostudiesfoundthatAd5-specificCD4TcellsaremoresusceptibletoHIVinfectionthanCD4Tcellsspecifictocertainothervectors,suchasCytomegalovirus[60]andCanarypox.[61] Bycomparison,aSciencearticlereportedthatChinahadapprovedCanSino'sEbolavaccinebasedonanAd5vector.ItwastestedinSierraLeone,whichhadhighHIVprevalence,makingitmorelikelyforsuchproblemstobedetected.CanSino'sCEOsaid"wehaven'tseenanythingwiththeEbolavaccine"andspeculatedthatHIVsusceptibilitymightbelimitedtoAd5vaccineswhichproducedHIVproteins.InresearchreportedinTheLancetinMay,thecompany'sresearchersacknowledgedthepossibility,calledit"controversial"andsaidtheywouldwatchforitinthecompany'sCOVID-19vaccinecandidate'strials.[46][50]ItisnotknowntowhatextentLGBTdiscriminationinSierraLeonecouldhavecontributedtomaskingapossiblecausallinkintheEbolavaccinetrial;whiletheSteptrialenrolledmainlyhomosexualandbisexualmen,thePhambilitrialenrolledmainlyheterosexualmenandwomenandstillfoundanapparentconnection.[citationneeded] Seealso[edit] Medicineportal Virusesportal VA(viralassociated)RNA References[edit] 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