Memory - Wikipedia

Memory is the faculty of the mind by which data or information is encoded, stored, and retrieved when needed. It is the retention of information over time ... Memory FromWikipedia,thefreeencyclopedia Jumptonavigation Jumptosearch Facultyofmindtostoreandretrievedata Thisarticleisabouthumanmemory.Forotheruses,seeMemory(disambiguation). Overviewoftheformsandfunctionsofmemory. Neuropsychology Topics Brainregions Clinicalneuropsychology Cognitiveneuropsychology Cognitiveneuroscience Dementia Humanbrain Neuroanatomy Neurophysiology Neuropsychologicalassessment Neuropsychologicalrehabilitation Traumaticbraininjury Brainfunctions Arousal Attention Consciousness Decisionmaking Executivefunctions Naturallanguage Learning Memory Motorcoordination Perception Planning Problemsolving Thought People AlanBaddeley ArthurL.Benton DavidBohm AntonioDamasio PhineasGage NormanGeschwind ElkhononGoldberg PatriciaGoldman-Rakic DonaldO.Hebb KennethHeilman EricKandel EdithKaplan MurielLezak BenjaminLibet RodolfoLlinás AlexanderLuria BrendaMilner KarlH.Pribram PaskoRakic OliverSacks MarkRosenzweig RogerW.Sperry Hans-LukasTeuber HenryMolaison("H.M.",patient) K.C.(patient) Tests BentonVisualRetentionTest ContinuousPerformanceTask Halstead-ReitanNeuropsychologicalBattery HaylingandBrixtontests LexicalDecisionTask Luria-Nebraskaneuropsychologicalbattery Mini–MentalStateExamination Rey–Osterriethcomplexfigure StroopTest WechslerAdultIntelligenceScale WechslerMemoryScale WisconsinCardSortingTask  Psychologyportal  Philosophyportal  Medicineportal vte Cognitivepsychology Perception Visualperception Objectrecognition Facerecognition Patternrecognition Attention Memory Agingandmemory Emotionalmemory Learning Long-termmemory Metacognition Language Metalanguage Thinking Cognition Concept Reasoning Decisionmaking Problemsolving Numericalcognition Numerosityadaptationeffect Approximatenumbersystem Parallelindividuationsystem vte Memoryisthefacultyofthemindbywhichdataorinformationisencoded,stored,andretrievedwhenneeded.Itistheretentionofinformationovertimeforthepurposeofinfluencingfutureaction.[1]Ifpasteventscouldnotberemembered,itwouldbeimpossibleforlanguage,relationships,orpersonalidentitytodevelop.[2]Memorylossisusuallydescribedasforgetfulnessoramnesia.[3][4][5][6][7][8] Memoryisoftenunderstoodasaninformationalprocessingsystemwithexplicitandimplicitfunctioningthatismadeupofasensoryprocessor,short-term(orworking)memory,andlong-termmemory.[9]Thiscanberelatedtotheneuron. Thesensoryprocessorallowsinformationfromtheoutsideworldtobesensedintheformofchemicalandphysicalstimuliandattendedtovariouslevelsoffocusandintent.Workingmemoryservesasanencodingandretrievalprocessor.Informationintheformofstimuliisencodedinaccordancewithexplicitorimplicitfunctionsbytheworkingmemoryprocessor.Theworkingmemoryalsoretrievesinformationfrompreviouslystoredmaterial.Finally,thefunctionoflong-termmemoryistostoredatathroughvariouscategoricalmodelsorsystems.[9] Declarative,orexplicit,memoryistheconsciousstorageandrecollectionofdata.[10]Underdeclarativememoryresidessemanticandepisodicmemory.Semanticmemoryreferstomemorythatisencodedwithspecificmeaning,[2]whileepisodicmemoryreferstoinformationthatisencodedalongaspatialandtemporalplane.[11][12][13]Declarativememoryisusuallytheprimaryprocessthoughtofwhenreferencingmemory.[2]Non-declarative,orimplicit,memoryistheunconsciousstorageandrecollectionofinformation.[14]Anexampleofanon-declarativeprocesswouldbetheunconsciouslearningorretrievalofinformationbywayofproceduralmemory,oraprimingphenomenon.[2][14][15]Primingistheprocessofsubliminallyarousingspecificresponsesfrommemoryandshowsthatnotallmemoryisconsciouslyactivated,[15]whereasproceduralmemoryistheslowandgraduallearningofskillsthatoftenoccurswithoutconsciousattentiontolearning.[2][14] Memoryisnotaperfectprocessor,andisaffectedbymanyfactors.Thewaysbywhichinformationisencoded,stored,andretrievedcanallbecorrupted.Pain,forexample,hasbeenidentifiedasaphysicalconditionthatimpairsmemory,andhasbeennotedinanimalmodelsaswellaschronicpainpatients.[16][17][18][19]Theamountofattentiongivennewstimulicandiminishtheamountofinformationthatbecomesencodedforstorage.[2]Also,thestorageprocesscanbecomecorruptedbyphysicaldamagetoareasofthebrainthatareassociatedwithmemorystorage,suchasthehippocampus.[20][21]Finally,theretrievalofinformationfromlong-termmemorycanbedisruptedbecauseofdecaywithinlong-termmemory.[2]Normalfunctioning,decayovertime,andbraindamageallaffecttheaccuracyandcapacityofthememory.[22][23] Contents 1Sensorymemory 2Short-termmemory 3Long-termmemory 3.1Multi-storemodel 3.2Workingmemory 4Types 4.1Byinformationtype 4.1.1Declarative 4.1.2Procedural 4.2Bytemporaldirection 5Studytechniques 5.1Toassessinfants 5.2Toassesschildrenandolderadults 6Failures 7Physiology 8Cognitiveneuroscience 9Genetics 9.1Geneticunderpinnings 9.2DNAmethylationanddemethylation 9.3Epigenetics 9.4RoleofDNATopoisomeraseIIBetainlearningandmemory 10Ininfancy 11Aging 12Disorders 13Influencingfactors 14Stress 15Sleep 16Constructionforgeneralmanipulation 17Improving 18Inplants 19Seealso 20Notes 20.1Sources 21Furtherreading 22Externallinks Sensorymemory[edit] Mainarticle:Sensorymemory Sensorymemoryholdsinformation,derivedfromthesenses,lessthanonesecondafteranitemisperceived.Theabilitytolookatanitemandrememberwhatitlookedlikewithjustasplitsecondofobservation,ormemorization,istheexampleofsensorymemory.Itisoutofcognitivecontrolandisanautomaticresponse.Withveryshortpresentations,participantsoftenreportthattheyseemto"see"morethantheycanactuallyreport.ThefirstpreciseexperimentsexploringthisformofsensorymemorywereconductedbyGeorgeSperling(1963)[24]usingthe"partialreportparadigm."Subjectswerepresentedwithagridof12letters,arrangedintothreerowsoffour.Afterabriefpresentation,subjectswerethenplayedeitherahigh,mediumorlowtone,cuingthemwhichoftherowstoreport.Basedonthesepartialreportexperiments,Sperlingwasabletoshowthatthecapacityofsensorymemorywasapproximately12items,butthatitdegradedveryquickly(withinafewhundredmilliseconds).Becausethisformofmemorydegradessoquickly,participantswouldseethedisplaybutbeunabletoreportalloftheitems(12inthe"wholereport"procedure)beforetheydecayed.Thistypeofmemorycannotbeprolongedviarehearsal. Threetypesofsensorymemoriesexist.Iconicmemoryisafastdecayingstoreofvisualinformation,atypeofsensorymemorythatbrieflystoresanimagethathasbeenperceivedforasmallduration.Echoicmemoryisafastdecayingstoreofauditoryinformation,alsoasensorymemorythatbrieflystoressoundsthathavebeenperceivedforshortdurations.[25][26]Hapticmemoryisatypeofsensorymemorythatrepresentsadatabasefortouchstimuli. Short-termmemory[edit] Mainarticle:Short-termmemory Short-termmemoryisalsoknownasworkingmemory.Short-termmemoryallowsrecallforaperiodofseveralsecondstoaminutewithoutrehearsal.Itscapacity,however,isverylimited.In1956,GeorgeA.Miller(1920–2012),whenworkingatBellLaboratories,conductedexperimentsshowingthatthestoreofshort-termmemorywas7±2items.(Hence,thetitleofhisfamouspaper,"TheMagicalNumber7±2.")Modernperspectivesestimatethecapacityofshort-termmemorytobelower,typicallyontheorderof4–5items,[27]orargueforamoreflexiblelimitbasedoninformationinsteadofitems.[28]Memorycapacitycanbeincreasedthroughaprocesscalledchunking.[29]Forexample,inrecallingaten-digittelephonenumber,apersoncouldchunkthedigitsintothreegroups:first,theareacode(suchas123),thenathree-digitchunk(456),and,last,afour-digitchunk(7890).Thismethodofrememberingtelephonenumbersisfarmoreeffectivethanattemptingtorememberastringof10digits;thisisbecauseweareabletochunktheinformationintomeaningfulgroupsofnumbers.Thisisreflectedinsomecountries'tendenciestodisplaytelephonenumbersasseveralchunksoftwotofournumbers. Short-termmemoryisbelievedtorelymostlyonanacousticcodeforstoringinformation,andtoalesserextentonavisualcode.Conrad(1964)[30]foundthattestsubjectshadmoredifficultyrecallingcollectionsoflettersthatwereacousticallysimilar,e.g.,E,P,D.Confusionwithrecallingacousticallysimilarlettersratherthanvisuallysimilarlettersimpliesthattheletterswereencodedacoustically.Conrad's(1964)study,however,dealswiththeencodingofwrittentext;thus,whilememoryofwrittenlanguagemayrelyonacousticcomponents,generalizationstoallformsofmemorycannotbemade. Long-termmemory[edit] Mainarticle:Long-termmemory OlinLeviWarner,Memory(1896).LibraryofCongressThomasJeffersonBuilding,Washington,D.C. Thestorageinsensorymemoryandshort-termmemorygenerallyhasastrictlylimitedcapacityandduration,whichmeansthatinformationisnotretainedindefinitely.Bycontrast,whilethetotalcapacityoflong-termmemoryhasyettobeestablished,itcanstoremuchlargerquantitiesofinformation.Furthermore,itcanstorethisinformationforamuchlongerduration,potentiallyforawholelifespan.Forexample,givenarandomseven-digitnumber,onemayrememberitforonlyafewsecondsbeforeforgetting,suggestingitwasstoredinshort-termmemory.Ontheotherhand,onecanremembertelephonenumbersformanyyearsthroughrepetition;thisinformationissaidtobestoredinlong-termmemory. Whileshort-termmemoryencodesinformationacoustically,long-termmemoryencodesitsemantically:Baddeley(1966)[31]discoveredthat,after20minutes,testsubjectshadthemostdifficultyrecallingacollectionofwordsthathadsimilarmeanings(e.g.big,large,great,huge)long-term.Anotherpartoflong-termmemoryisepisodicmemory,"whichattemptstocaptureinformationsuchas'what','when'and'where'".[32]Withepisodicmemory,individualsareabletorecallspecificeventssuchasbirthdaypartiesandweddings. Short-termmemoryissupportedbytransientpatternsofneuronalcommunication,dependentonregionsofthefrontallobe(especiallydorsolateralprefrontalcortex)andtheparietallobe.Long-termmemory,ontheotherhand,ismaintainedbymorestableandpermanentchangesinneuralconnectionswidelyspreadthroughoutthebrain.Thehippocampusisessential(forlearningnewinformation)totheconsolidationofinformationfromshort-termtolong-termmemory,althoughitdoesnotseemtostoreinformationitself.Itwasthoughtthatwithoutthehippocampusnewmemorieswereunabletobestoredintolong-termmemoryandthattherewouldbeaveryshortattentionspan,asfirstgleanedfrompatientHenryMolaison[33][34]afterwhatwasthoughttobethefullremovalofbothhishippocampi.Morerecentexaminationofhisbrain,post-mortem,showsthatthehippocampuswasmoreintactthanfirstthought,throwingtheoriesdrawnfromtheinitialdataintoquestion.Thehippocampusmaybeinvolvedinchangingneuralconnectionsforaperiodofthreemonthsormoreaftertheinitiallearning. Researchhassuggestedthatlong-termmemorystorageinhumansmaybemaintainedbyDNAmethylation,[35]andthe'prion'gene.[36][37] Furtherresearchinvestigatedthemolecularbasisforlong-termmemory.By2015ithadbecomeclearthatlong-termmemoryrequiresgenetranscriptionactivationanddenovoproteinsynthesis.[38]Long-termmemoryformationdependsonboththeactivationofmemorypromotinggenesandtheinhibitionofmemorysuppressorgenes,andDNAmethylation/DNAdemethylationwasfoundtobeamajormechanismforachievingthisdualregulation.[39] Ratswithanew,stronglong-termmemoryduetocontextualfearconditioninghavereducedexpressionofabout1,000genesandincreasedexpressionofabout500genesinthehippocampus24hoursaftertraining,thusexhibitingmodifiedexpressionof9.17%oftherathippocampalgenome.Reducedgeneexpressionswereassociatedwithmethylationsofthosegenes.[40] Considerablefurtherresearchintolong-termmemoryhasilluminatedthemolecularmechanismsbywhichmethylationsareestablishedorremoved,asreviewedin2022.[41]Thesemechanismsinclude,forinstance,signal-responsiveTOP2B-induceddouble-strandbreaksinimmediateearlygenes.AlsothemessengerRNAsofmanygenesthathadbeensubjectedtomethylation-controlledincreasesordecreasesaretransportedbyneuralgranules(messengerRNP)tothedendriticspines.AttheselocationsthemessengerRNAscanbetranslatedintotheproteinsthatcontrolsignalingatneuronalsynapses.[41] Multi-storemodel[edit] Seealso:Memoryconsolidation Multi-storemodel Themulti-storemodel(alsoknownasAtkinson–Shiffrinmemorymodel)wasfirstdescribedin1968byAtkinsonandShiffrin. Themulti-storemodelhasbeencriticisedforbeingtoosimplistic.Forinstance,long-termmemoryisbelievedtobeactuallymadeupofmultiplesubcomponents,suchasepisodicandproceduralmemory.Italsoproposesthatrehearsalistheonlymechanismbywhichinformationeventuallyreacheslong-termstorage,butevidenceshowsuscapableofrememberingthingswithoutrehearsal. Themodelalsoshowsallthememorystoresasbeingasingleunitwhereasresearchintothisshowsdifferently.Forexample,short-termmemorycanbebrokenupintodifferentunitssuchasvisualinformationandacousticinformation.InastudybyZlonogaandGerber(1986),patient'KF'demonstratedcertaindeviationsfromtheAtkinson–Shiffrinmodel.PatientKFwasbraindamaged,displayingdifficultiesregardingshort-termmemory.Recognitionofsoundssuchasspokennumbers,letters,wordsandeasilyidentifiablenoises(suchasdoorbellsandcatsmeowing)wereallimpacted.Visualshort-termmemorywasunaffected,suggestingadichotomybetweenvisualandaudialmemory.[42] Workingmemory[edit] Mainarticle:Workingmemory Theworkingmemorymodel In1974BaddeleyandHitchproposeda"workingmemorymodel"thatreplacedthegeneralconceptofshort-termmemorywithanactivemaintenanceofinformationintheshort-termstorage.Inthismodel,workingmemoryconsistsofthreebasicstores:thecentralexecutive,thephonologicalloopandthevisuo-spatialsketchpad.In2000thismodelwasexpandedwiththemultimodalepisodicbuffer(Baddeley'smodelofworkingmemory).[43] Thecentralexecutiveessentiallyactsasanattentionsensorystore.Itchannelsinformationtothethreecomponentprocesses:thephonologicalloop,thevisuo-spatialsketchpad,andtheepisodicbuffer. Thephonologicalloopstoresauditoryinformationbysilentlyrehearsingsoundsorwordsinacontinuousloop:thearticulatoryprocess(forexampletherepetitionofatelephonenumberoverandoveragain).Ashortlistofdataiseasiertoremember.Thephonologicalloopisoccasionallydisrupted.Irrelevantspeechorbackgroundnoisecanimpedethephonologicalloop.Articulatorysuppressioncanalsoconfuseencodingandwordsthatsoundsimilarcanbeswitchedormisrememberedthroughthephonologicalsimilarityeffect.thephonologicalloopalsohasalimittohowmuchitcanholdatoncewhichmeansthatitiseasiertorememberalotofshortwordsratherthanalotoflongwords,accordingtothewordlengtheffect. Thevisuospatialsketchpadstoresvisualandspatialinformation.Itisengagedwhenperformingspatialtasks(suchasjudgingdistances)orvisualones(suchascountingthewindowsonahouseorimaginingimages).ThosewithAphantasiawillnotbeabletoengagethevisuospatialsketchpad. Theepisodicbufferisdedicatedtolinkinginformationacrossdomainstoformintegratedunitsofvisual,spatial,andverbalinformationandchronologicalordering(e.g.,thememoryofastoryoramoviescene).Theepisodicbufferisalsoassumedtohavelinkstolong-termmemoryandsemanticalmeaning. Theworkingmemorymodelexplainsmanypracticalobservations,suchaswhyitiseasiertodotwodifferenttasks(oneverbalandonevisual)thantwosimilartasks(e.g.,twovisual),andtheaforementionedword-lengtheffect.Workingmemoryisalsothepremiseforwhatallowsustodoeverydayactivitiesinvolvingthought.Itisthesectionofmemorywherewecarryoutthoughtprocessesandusethemtolearnandreasonabouttopics.[43] Types[edit] Researchersdistinguishbetweenrecognitionandrecallmemory.Recognitionmemorytasksrequireindividualstoindicatewhethertheyhaveencounteredastimulus(suchasapictureoraword)before.Recallmemorytasksrequireparticipantstoretrievepreviouslylearnedinformation.Forexample,individualsmightbeaskedtoproduceaseriesofactionstheyhaveseenbeforeortosayalistofwordstheyhaveheardbefore. Byinformationtype[edit] Topographicalmemoryinvolvestheabilitytoorientoneselfinspace,torecognizeandfollowanitinerary,ortorecognizefamiliarplaces.[44]Gettinglostwhentravelingaloneisanexampleofthefailureoftopographicmemory.[45] Flashbulbmemoriesareclearepisodicmemoriesofuniqueandhighlyemotionalevents.[46]PeoplerememberingwheretheywereorwhattheyweredoingwhentheyfirstheardthenewsofPresidentKennedy'sassassination,[47]theSydneySiegeorof9/11areexamplesofflashbulbmemories. Anderson(1976)[48]divideslong-termmemoryintodeclarative(explicit)andprocedural(implicit)memories. Declarative[edit] Mainarticle:Declarativememory Declarativememoryrequiresconsciousrecall,inthatsomeconsciousprocessmustcallbacktheinformation.Itissometimescalledexplicitmemory,sinceitconsistsofinformationthatisexplicitlystoredandretrieved.Declarativememorycanbefurthersub-dividedintosemanticmemory,concerningprinciplesandfactstakenindependentofcontext;andepisodicmemory,concerninginformationspecifictoaparticularcontext,suchasatimeandplace.Semanticmemoryallowstheencodingofabstractknowledgeabouttheworld,suchas"ParisisthecapitalofFrance".Episodicmemory,ontheotherhand,isusedformorepersonalmemories,suchasthesensations,emotions,andpersonalassociationsofaparticularplaceortime.Episodicmemoriesoftenreflectthe"firsts"inlifesuchasafirstkiss,firstdayofschoolorfirsttimewinningachampionship.Thesearekeyeventsinone'slifethatcanberememberedclearly. Researchsuggeststhatdeclarativememoryissupportedbyseveralfunctionsofthemedialtemporallobesystemwhichincludesthehippocampus.[49]Autobiographicalmemory–memoryforparticulareventswithinone'sownlife–isgenerallyviewedaseitherequivalentto,orasubsetof,episodicmemory.Visualmemoryispartofmemorypreservingsomecharacteristicsofoursensespertainingtovisualexperience.Oneisabletoplaceinmemoryinformationthatresemblesobjects,places,animalsorpeopleinsortofamentalimage.Visualmemorycanresultinpriminganditisassumedsomekindofperceptualrepresentationalsystemunderliesthisphenomenon.[49] Procedural[edit] Incontrast,proceduralmemory(orimplicitmemory)isnotbasedontheconsciousrecallofinformation,butonimplicitlearning.Itcanbestbesummarizedasrememberinghowtodosomething.Proceduralmemoryisprimarilyusedinlearningmotorskillsandcanbeconsideredasubsetofimplicitmemory.Itisrevealedwhenonedoesbetterinagiventaskdueonlytorepetition–nonewexplicitmemorieshavebeenformed,butoneisunconsciouslyaccessingaspectsofthosepreviousexperiences.Proceduralmemoryinvolvedinmotorlearningdependsonthecerebellumandbasalganglia.[50] Acharacteristicofproceduralmemoryisthatthethingsrememberedareautomaticallytranslatedintoactions,andthussometimesdifficulttodescribe.Someexamplesofproceduralmemoryincludetheabilitytorideabikeortieshoelaces.[51] Bytemporaldirection[edit] Anothermajorwaytodistinguishdifferentmemoryfunctionsiswhetherthecontenttoberememberedisinthepast,retrospectivememory,orinthefuture,prospectivememory.JohnMeachamintroducedthisdistinctioninapaperpresentedatthe1975AmericanPsychologicalAssociationannualmeetingandsubsequentlyincludedbyUlricNeisserinhis1982editedvolume,MemoryObserved:RememberinginNaturalContexts.[52][53]Thus,retrospectivememoryasacategoryincludessemantic,episodicandautobiographicalmemory.Incontrast,prospectivememoryismemoryforfutureintentions,orrememberingtoremember(Winograd,1988).Prospectivememorycanbefurtherbrokendownintoevent-andtime-basedprospectiveremembering.Time-basedprospectivememoriesaretriggeredbyatime-cue,suchasgoingtothedoctor(action)at4pm(cue).Event-basedprospectivememoriesareintentionstriggeredbycues,suchasrememberingtopostaletter(action)afterseeingamailbox(cue).Cuesdonotneedtoberelatedtotheaction(asthemailbox/letterexample),andlists,sticky-notes,knottedhandkerchiefs,orstringaroundthefingerallexemplifycuesthatpeopleuseasstrategiestoenhanceprospectivememory. Studytechniques[edit] Toassessinfants[edit] Infantsdonothavethelanguageabilitytoreportontheirmemoriesandsoverbalreportscannotbeusedtoassessveryyoungchildren'smemory.Throughouttheyears,however,researchershaveadaptedanddevelopedanumberofmeasuresforassessingbothinfants'recognitionmemoryandtheirrecallmemory.Habituationandoperantconditioningtechniqueshavebeenusedtoassessinfants'recognitionmemoryandthedeferredandelicitedimitationtechniqueshavebeenusedtoassessinfants'recallmemory. Techniquesusedtoassessinfants'recognitionmemoryincludethefollowing: Visualpairedcomparisonprocedure(reliesonhabituation):infantsarefirstpresentedwithpairsofvisualstimuli,suchastwoblack-and-whitephotosofhumanfaces,forafixedamountoftime;then,afterbeingfamiliarizedwiththetwophotos,theyarepresentedwiththe"familiar"photoandanewphoto.Thetimespentlookingateachphotoisrecorded.Lookinglongeratthenewphotoindicatesthattheyrememberthe"familiar"one.Studiesusingthisprocedurehavefoundthat5-to6-month-oldscanretaininformationforaslongasfourteendays.[54] Operantconditioningtechnique:infantsareplacedinacribandaribbonthatisconnectedtoamobileoverheadistiedtooneoftheirfeet.Infantsnoticethatwhentheykicktheirfootthemobilemoves–therateofkickingincreasesdramaticallywithinminutes.Studiesusingthistechniquehaverevealedthatinfants'memorysubstantiallyimprovesoverthefirst18-months.Whereas2-to3-month-oldscanretainanoperantresponse(suchasactivatingthemobilebykickingtheirfoot)foraweek,6-month-oldscanretainitfortwoweeks,and18-month-oldscanretainasimilaroperantresponseforaslongas13weeks.[55][56][57] Techniquesusedtoassessinfants'recallmemoryincludethefollowing: Deferredimitationtechnique:anexperimentershowsinfantsauniquesequenceofactions(suchasusingasticktopushabuttononabox)andthen,afteradelay,askstheinfantstoimitatetheactions.Studiesusingdeferredimitationhaveshownthat14-month-olds'memoriesforthesequenceofactionscanlastforaslongasfourmonths.[58] Elicitedimitationtechnique:isverysimilartothedeferredimitationtechnique;thedifferenceisthatinfantsareallowedtoimitatetheactionsbeforethedelay.Studiesusingtheelicitedimitationtechniquehaveshownthat20-month-oldscanrecalltheactionsequencestwelvemonthslater.[59][60] Toassesschildrenandolderadults[edit] Researchersuseavarietyoftaskstoassessolderchildrenandadults'memory.Someexamplesare: Pairedassociatelearning–whenonelearnstoassociateonespecificwordwithanother.Forexample,whengivenawordsuchas"safe"onemustlearntosayanotherspecificword,suchas"green".Thisisstimulusandresponse.[61][62] Freerecall–duringthistaskasubjectwouldbeaskedtostudyalistofwordsandthenlatertheywillbeaskedtorecallorwritedownasmanywordsthattheycanremember,similartofreeresponsequestions.[63]Earlieritemsareaffectedbyretroactiveinterference(RI),whichmeansthelongerthelist,thegreatertheinterference,andthelesslikelihoodthattheyarerecalled.Ontheotherhand,itemsthathavebeenpresentedlastlysufferlittleRI,butsufferagreatdealfromproactiveinterference(PI),whichmeansthelongerthedelayinrecall,themorelikelythattheitemswillbelost.[64] Cuedrecall–oneisgivenasignificanthintstohelpretrieveinformationthathasbeenpreviouslyencodedintotheperson'smemory;typicallythiscaninvolveawordrelatingtotheinformationbeingaskedtoremember.[65]Thisissimilartofillintheblankassessmentsusedinclassrooms. Recognition–subjectsareaskedtorememberalistofwordsorpictures,afterwhichpointtheyareaskedtoidentifythepreviouslypresentedwordsorpicturesfromamongalistofalternativesthatwerenotpresentedintheoriginallist.[66]Thisissimilartomultiplechoiceassessments. Detectionparadigm–individualsareshownanumberofobjectsandcolorsamplesduringacertainperiodoftime.Theyarethentestedontheirvisualabilitytorememberasmuchastheycanbylookingattestersandpointingoutwhetherthetestersaresimilartothesample,orifanychangeispresent. Savingsmethod–comparesthespeedoforiginallylearningtothespeedofrelearningit.Theamountoftimesavedmeasuresmemory.[67] Implicit-memorytasks–informationisdrawnfrommemorywithoutconsciousrealization. Failures[edit] Seealso:Eyewitnessmemory Thegardenofoblivion,illustrationbyEphraimMosesLilien. Transience–memoriesdegradewiththepassingoftime.Thisoccursinthestoragestageofmemory,aftertheinformationhasbeenstoredandbeforeitisretrieved.Thiscanhappeninsensory,short-term,andlong-termstorage.Itfollowsageneralpatternwheretheinformationisrapidlyforgottenduringthefirstcoupleofdaysoryears,followedbysmalllossesinlaterdaysoryears. Absent-mindedness–Memoryfailureduetothelackofattention.Attentionplaysakeyroleinstoringinformationintolong-termmemory;withoutproperattention,theinformationmightnotbestored,makingitimpossibletoberetrievedlater. Physiology[edit] Brainareasinvolvedintheneuroanatomyofmemorysuchasthehippocampus,theamygdala,thestriatum,orthemammillarybodiesarethoughttobeinvolvedinspecifictypesofmemory.Forexample,thehippocampusisbelievedtobeinvolvedinspatiallearninganddeclarativelearning,whiletheamygdalaisthoughttobeinvolvedinemotionalmemory.[68] Damagetocertainareasinpatientsandanimalmodelsandsubsequentmemorydeficitsisaprimarysourceofinformation.However,ratherthanimplicatingaspecificarea,itcouldbethatdamagetoadjacentareas,ortoapathwaytravelingthroughtheareaisactuallyresponsiblefortheobserveddeficit.Further,itisnotsufficienttodescribememory,anditscounterpart,learning,assolelydependentonspecificbrainregions.Learningandmemoryareusuallyattributedtochangesinneuronalsynapses,thoughttobemediatedbylong-termpotentiationandlong-termdepression. Ingeneral,themoreemotionallychargedaneventorexperienceis,thebetteritisremembered;thisphenomenonisknownasthememoryenhancementeffect.Patientswithamygdaladamage,however,donotshowamemoryenhancementeffect.[69][70] Hebbdistinguishedbetweenshort-termandlong-termmemory.Hepostulatedthatanymemorythatstayedinshort-termstorageforalongenoughtimewouldbeconsolidatedintoalong-termmemory.Laterresearchshowedthistobefalse.Researchhasshownthatdirectinjectionsofcortisolorepinephrinehelpthestorageofrecentexperiences.Thisisalsotrueforstimulationoftheamygdala.Thisprovesthatexcitementenhancesmemorybythestimulationofhormonesthataffecttheamygdala.Excessiveorprolongedstress(withprolongedcortisol)mayhurtmemorystorage.Patientswithamygdalardamagearenomorelikelytorememberemotionallychargedwordsthannonemotionallychargedones.Thehippocampusisimportantforexplicitmemory.Thehippocampusisalsoimportantformemoryconsolidation.Thehippocampusreceivesinputfromdifferentpartsofthecortexandsendsitsoutputouttodifferentpartsofthebrainalso.Theinputcomesfromsecondaryandtertiarysensoryareasthathaveprocessedtheinformationalotalready.Hippocampaldamagemayalsocausememorylossandproblemswithmemorystorage.[71]Thismemorylossincludesretrogradeamnesiawhichisthelossofmemoryforeventsthatoccurredshortlybeforethetimeofbraindamage.[67] Cognitiveneuroscience[edit] Cognitiveneuroscientistsconsidermemoryastheretention,reactivation,andreconstructionoftheexperience-independentinternalrepresentation.Thetermofinternalrepresentationimpliesthatsuchadefinitionofmemorycontainstwocomponents:theexpressionofmemoryatthebehavioralorconsciouslevel,andtheunderpinningphysicalneuralchanges(Dudai2007).Thelattercomponentisalsocalledengramormemorytraces(Semon1904).Someneuroscientistsandpsychologistsmistakenlyequatetheconceptofengramandmemory,broadlyconceivingallpersistingafter-effectsofexperiencesasmemory;othersargueagainstthisnotionthatmemorydoesnotexistuntilitisrevealedinbehaviororthought(Moscovitch2007). Onequestionthatiscrucialincognitiveneuroscienceishowinformationandmentalexperiencesarecodedandrepresentedinthebrain.Scientistshavegainedmuchknowledgeabouttheneuronalcodesfromthestudiesofplasticity,butmostofsuchresearchhasbeenfocusedonsimplelearninginsimpleneuronalcircuits;itisconsiderablylessclearabouttheneuronalchangesinvolvedinmorecomplexexamplesofmemory,particularlydeclarativememorythatrequiresthestorageoffactsandevents(Byrne2007).Convergence-divergencezonesmightbetheneuralnetworkswherememoriesarestoredandretrieved.Consideringthatthereareseveralkindsofmemory,dependingontypesofrepresentedknowledge,underlyingmechanisms,processesfunctionsandmodesofacquisition,itislikelythatdifferentbrainareassupportdifferentmemorysystemsandthattheyareinmutualrelationshipsinneuronalnetworks:"componentsofmemoryrepresentationaredistributedwidelyacrossdifferentpartsofthebrainasmediatedbymultipleneocorticalcircuits".[72] Encoding.Encodingofworkingmemoryinvolvesthespikingofindividualneuronsinducedbysensoryinput,whichpersistsevenafterthesensoryinputdisappears(JensenandLisman2005;Fransenetal.2002).Encodingofepisodicmemoryinvolvespersistentchangesinmolecularstructuresthataltersynaptictransmissionbetweenneurons.Examplesofsuchstructuralchangesincludelong-termpotentiation(LTP)orspike-timing-dependentplasticity(STDP).Thepersistentspikinginworkingmemorycanenhancethesynapticandcellularchangesintheencodingofepisodicmemory(JensenandLisman2005). Workingmemory.Recentfunctionalimagingstudiesdetectedworkingmemorysignalsinbothmedialtemporallobe(MTL),abrainareastronglyassociatedwithlong-termmemory,andprefrontalcortex(Ranganathetal.2005),suggestingastrongrelationshipbetweenworkingmemoryandlong-termmemory.However,thesubstantiallymoreworkingmemorysignalsseenintheprefrontallobesuggestthatthisareaplaysamoreimportantroleinworkingmemorythanMTL(Suzuki2007). Consolidationandreconsolidation.Short-termmemory(STM)istemporaryandsubjecttodisruption,whilelong-termmemory(LTM),onceconsolidated,ispersistentandstable.ConsolidationofSTMintoLTMatthemolecularlevelpresumablyinvolvestwoprocesses:synapticconsolidationandsystemconsolidation.Theformerinvolvesaproteinsynthesisprocessinthemedialtemporallobe(MTL),whereasthelattertransformstheMTL-dependentmemoryintoanMTL-independentmemoryovermonthstoyears(Ledoux2007).Inrecentyears,suchtraditionalconsolidationdogmahasbeenre-evaluatedasaresultofthestudiesonreconsolidation.Thesestudiesshowedthatpreventionafterretrievalaffectssubsequentretrievalofthememory(Sara2000).Newstudieshaveshownthatpost-retrievaltreatmentwithproteinsynthesisinhibitorsandmanyothercompoundscanleadtoanamnesticstate(Nadeletal.2000b;Alberini2005;Dudai2006).Thesefindingsonreconsolidationfitwiththebehavioralevidencethatretrievedmemoryisnotacarboncopyoftheinitialexperiences,andmemoriesareupdatedduringretrieval. Genetics[edit] Seealso:Long-termpotentiationandEricKandel Studyofthegeneticsofhumanmemoryisinitsinfancythoughmanygeneshavebeeninvestigatedfortheirassociationtomemoryinhumansandnon-humananimals.AnotableinitialsuccesswastheassociationofAPOEwithmemorydysfunctioninAlzheimer'sdisease.Thesearchforgenesassociatedwithnormallyvaryingmemorycontinues.OneofthefirstcandidatesfornormalvariationinmemoryistheproteinKIBRA,[73][medicalcitationneeded]whichappearstobeassociatedwiththerateatwhichmaterialisforgottenoveradelayperiod.Therehasbeensomeevidencethatmemoriesarestoredinthenucleusofneurons.[74][medicalcitationneeded] Geneticunderpinnings[edit] Severalgenes,proteinsandenzymeshavebeenextensivelyresearchedfortheirassociationwithmemory.Long-termmemory,unlikeshort-termmemory,isdependentuponthesynthesisofnewproteins.[75]Thisoccurswithinthecellularbody,andconcernstheparticulartransmitters,receptors,andnewsynapsepathwaysthatreinforcethecommunicativestrengthbetweenneurons.Theproductionofnewproteinsdevotedtosynapsereinforcementistriggeredafterthereleaseofcertainsignalingsubstances(suchascalciumwithinhippocampalneurons)inthecell.Inthecaseofhippocampalcells,thisreleaseisdependentupontheexpulsionofmagnesium(abindingmolecule)thatisexpelledaftersignificantandrepetitivesynapticsignaling.ThetemporaryexpulsionofmagnesiumfreesNMDAreceptorstoreleasecalciuminthecell,asignalthatleadstogenetranscriptionandtheconstructionofreinforcingproteins.[76]Formoreinformation,seelong-termpotentiation(LTP). OneofthenewlysynthesizedproteinsinLTPisalsocriticalformaintaininglong-termmemory.ThisproteinisanautonomouslyactiveformoftheenzymeproteinkinaseC(PKC),knownasPKMζ.PKMζmaintainstheactivity-dependentenhancementofsynapticstrengthandinhibitingPKMζerasesestablishedlong-termmemories,withoutaffectingshort-termmemoryor,oncetheinhibitoriseliminated,theabilitytoencodeandstorenewlong-termmemoriesisrestored.Also,BDNFisimportantforthepersistenceoflong-termmemories.[77] Thelong-termstabilizationofsynapticchangesisalsodeterminedbyaparallelincreaseofpre-andpostsynapticstructuressuchasaxonalbouton,dendriticspineandpostsynapticdensity.[78] Onthemolecularlevel,anincreaseofthepostsynapticscaffoldingproteinsPSD-95andHOMER1chasbeenshowntocorrelatewiththestabilizationofsynapticenlargement.[78]ThecAMPresponseelement-bindingprotein(CREB)isatranscriptionfactorwhichisbelievedtobeimportantinconsolidatingshort-termtolong-termmemories,andwhichisbelievedtobedownregulatedinAlzheimer'sdisease.[79] DNAmethylationanddemethylation[edit] Ratsexposedtoanintenselearningeventmayretainalife-longmemoryoftheevent,evenafterasingletrainingsession.Thelong-termmemoryofsuchaneventappearstobeinitiallystoredinthehippocampus,butthisstorageistransient.Muchofthelong-termstorageofthememoryseemstotakeplaceintheanteriorcingulatecortex.[80]Whensuchanexposurewasexperimentallyapplied,morethan5,000differentlymethylatedDNAregionsappearedinthehippocampusneuronalgenomeoftheratsatoneandat24hoursaftertraining.[81]Thesealterationsinmethylationpatternoccurredatmanygenesthatweredown-regulated,oftenduetotheformationofnew5-methylcytosinesitesinCpGrichregionsofthegenome.Furthermore,manyothergeneswereupregulated,likelyoftenduetohypomethylation.Hypomethylationoftenresultsfromtheremovalofmethylgroupsfrompreviouslyexisting5-methylcytosinesinDNA.Demethylationiscarriedoutbyseveralproteinsactinginconcert,includingtheTETenzymesaswellasenzymesoftheDNAbaseexcisionrepairpathway(seeEpigeneticsinlearningandmemory).Thepatternofinducedandrepressedgenesinbrainneuronssubsequenttoanintenselearningeventlikelyprovidesthemolecularbasisforalong-termmemoryoftheevent. Epigenetics[edit] Mainarticle:Epigeneticsinlearningandmemory Studiesofthemolecularbasisformemoryformationindicatethatepigeneticmechanismsoperatinginbrainneuronsplayacentralroleindeterminingthiscapability.KeyepigeneticmechanismsinvolvedinmemoryincludethemethylationanddemethylationofneuronalDNA,aswellasmodificationsofhistoneproteinsincludingmethylations,acetylationsanddeacetylations. StimulationofbrainactivityinmemoryformationisoftenaccompaniedbythegenerationofdamageinneuronalDNAthatisfollowedbyrepairassociatedwithpersistentepigeneticalterations.InparticulartheDNArepairprocessesofnon-homologousendjoiningandbaseexcisionrepairareemployedinmemoryformation.[citationneeded] RoleofDNATopoisomeraseIIBetainlearningandmemory[edit] Duringanewlearningexperience,asetofgenesisrapidlyexpressedinthebrain.Thisinducedgeneexpressionisconsideredtobeessentialforprocessingtheinformationbeinglearned.Suchgenesarereferredtoasimmediateearlygenes(IEGs).DNATopoisomeraseIIBeta(TOP2B)activityisessentialfortheexpressionofIEGsinatypeoflearningexperienceinmicetermedassociativefearmemory.[82]SuchalearningexperienceappearstorapidlytriggerTOP2Btoinducedouble-strandbreaksinthepromoterDNAofIEGgenesthatfunctioninneuroplasticity.RepairoftheseinducedbreaksisassociatedwithDNAdemethylationofIEGgenepromotersallowingimmediateexpressionoftheseIEGgenes.[82] RegulatorysequenceinapromoteratatranscriptionstartsitewithapausedRNApolymeraseandaTOP2B-induceddouble-strandbreak Thedouble-strandbreaksthatareinducedduringalearningexperiencearenotimmediatelyrepaired.About600regulatorysequencesinpromotersandabout800regulatorysequencesinenhancersappeartodependondoublestrandbreaksinitiatedbytopoisomerase2-beta(TOP2B)foractivation.[83][84]Theinductionofparticulardouble-strandbreaksarespecificwithrespecttotheirinducingsignal.Whenneuronsareactivatedinvitro,just22ofTOP2B-induceddouble-strandbreaksoccurintheirgenomes.[85] SuchTOP2B-induceddouble-strandbreaksareaccompaniedbyatleastfourenzymesofthenon-homologousendjoining(NHEJ)DNArepairpathway(DNA-PKcs,KU70,KU80,andDNALIGASEIV)(seeFigure).Theseenzymesrepairthedouble-strandbreakswithinabout15minutestotwohours.[85][86]Thedouble-strandbreaksinthepromoterarethusassociatedwithTOP2Bandatleastthesefourrepairenzymes.Theseproteinsarepresentsimultaneouslyonasinglepromoternucleosome(thereareabout147nucleotidesintheDNAsequencewrappedaroundasinglenucleosome)locatednearthetranscriptionstartsiteoftheirtargetgene.[86] Brainregionsinvolvedinmemoryformationincludingmedialprefrontalcortex(mPFC) Thedouble-strandbreakintroducedbyTOP2BapparentlyfreesthepartofthepromoteratanRNApolymerase-boundtranscriptionstartsitetophysicallymovetoitsassociatedenhancer(seeregulatorysequence).Thisallowstheenhancer,withitsboundtranscriptionfactorsandmediatorproteins,todirectlyinteractwiththeRNApolymerasepausedatthetranscriptionstartsitetostarttranscription.[85][87] Contextualfearconditioninginthemousecausesthemousetohavealong-termmemoryandfearofthelocationinwhichitoccurred.ContextualfearconditioningcauseshundredsofDSBsinmousebrainmedialprefrontalcortex(mPFC)andhippocampusneurons(seeFigure:Brainregionsinvolvedinmemoryformation).TheseDSBspredominatelyactivategenesinvolvedinsynapticprocesses,thatareimportantforlearningandmemory.[88] Ininfancy[edit] Fortheinabilityofadultstoretrieveearlymemories,seeChildhoodamnesia. Upuntilthemid-1980sitwasassumedthatinfantscouldnotencode,retain,andretrieveinformation.[89]Agrowingbodyofresearchnowindicatesthatinfantsasyoungas6-monthscanrecallinformationaftera24-hourdelay.[90]Furthermore,researchhasrevealedthatasinfantsgrowoldertheycanstoreinformationforlongerperiodsoftime;6-month-oldscanrecallinformationaftera24-hourperiod,9-month-oldsafteruptofiveweeks,and20-month-oldsafteraslongastwelvemonths.[91]Inaddition,studieshaveshownthatwithage,infantscanstoreinformationfaster.Whereas14-month-oldscanrecallathree-stepsequenceafterbeingexposedtoitonce,6-month-oldsneedapproximatelysixexposuresinordertobeabletorememberit.[58][90] Although6-month-oldscanrecallinformationovertheshort-term,theyhavedifficultyrecallingthetemporalorderofinformation.Itisonlyby9monthsofagethatinfantscanrecalltheactionsofatwo-stepsequenceinthecorrecttemporalorder–thatis,recallingstep1andthenstep2.[92][93]Inotherwords,whenaskedtoimitateatwo-stepactionsequence(suchasputtingatoycarinthebaseandpushingintheplungertomakethetoyrolltotheotherend),9-month-oldstendtoimitatetheactionsofthesequenceinthecorrectorder(step1andthenstep2).Youngerinfants(6-month-olds)canonlyrecallonestepofatwo-stepsequence.[90]Researchershavesuggestedthattheseagedifferencesareprobablyduetothefactthatthedentategyrusofthehippocampusandthefrontalcomponentsoftheneuralnetworkarenotfullydevelopedattheageof6-months.[59][94][95] Infact,theterm'infantileamnesia'referstothephenomenonofacceleratedforgettingduringinfancy.Importantly,infantileamnesiaisnotuniquetohumans,andpreclinicalresearch(usingrodentmodels)providesinsightintothepreciseneurobiologyofthisphenomenon.AreviewoftheliteraturefrombehavioralneuroscientistDrJeeHyunKimsuggeststhatacceleratedforgettingduringearlylifeisatleastpartlyduetorapidgrowthofthebrainduringthisperiod.[96] Aging[edit] Mainarticle:Memoryandaging Oneofthekeyconcernsofolderadultsistheexperienceofmemoryloss,especiallyasitisoneofthehallmarksymptomsofAlzheimer'sdisease.However,memorylossisqualitativelydifferentinnormalagingfromthekindofmemorylossassociatedwithadiagnosisofAlzheimer's(Budson&Price,2005).Researchhasrevealedthatindividuals'performanceonmemorytasksthatrelyonfrontalregionsdeclineswithage.Olderadultstendtoexhibitdeficitsontasksthatinvolveknowingthetemporalorderinwhichtheylearnedinformation;[97]sourcememorytasksthatrequirethemtorememberthespecificcircumstancesorcontextinwhichtheylearnedinformation;[98]andprospectivememorytasksthatinvolverememberingtoperformanactatafuturetime.Olderadultscanmanagetheirproblemswithprospectivememorybyusingappointmentbooks,forexample. Genetranscriptionprofilesweredeterminedforthehumanfrontalcortexofindividualsfromage26to106years.Numerousgeneswereidentifiedwithreducedexpressionafterage40,andespeciallyafterage70.[99]Genesthatplaycentralrolesinmemoryandlearningwereamongthoseshowingthemostsignificantreductionwithage.TherewasalsoamarkedincreaseinDNAdamage,likelyoxidativedamage,inthepromotersofthosegeneswithreducedexpression.ItwassuggestedthatDNAdamagemayreducetheexpressionofselectivelyvulnerablegenesinvolvedinmemoryandlearning.[99] Disorders[edit] Mainarticle:Memorydisorder Muchofthecurrentknowledgeofmemoryhascomefromstudyingmemorydisorders,particularlylossofmemory,knownasamnesia.Amnesiacanresultfromextensivedamageto:(a)theregionsofthemedialtemporallobe,suchasthehippocampus,dentategyrus,subiculum,amygdala,theparahippocampal,entorhinal,andperirhinalcortices[100]orthe(b)midlinediencephalicregion,specificallythedorsomedialnucleusofthethalamusandthemammillarybodiesofthehypothalamus.[101]Therearemanysortsofamnesia,andbystudyingtheirdifferentforms,ithasbecomepossibletoobserveapparentdefectsinindividualsub-systemsofthebrain'smemorysystems,andthushypothesizetheirfunctioninthenormallyworkingbrain.OtherneurologicaldisorderssuchasAlzheimer'sdiseaseandParkinson'sdisease[102][better source needed]canalsoaffectmemoryandcognition. Hyperthymesia,orhyperthymesicsyndrome,isadisorderthataffectsanindividual'sautobiographicalmemory,essentiallymeaningthattheycannotforgetsmalldetailsthatotherwisewouldnotbestored.[103][medicalcitationneeded]Korsakoff'ssyndrome,alsoknownasKorsakoff'spsychosis,amnesic-confabulatorysyndrome,isanorganicbraindiseasethatadverselyaffectsmemorybywidespreadlossorshrinkageofneuronswithintheprefrontalcortex.[67] Whilenotadisorder,acommontemporaryfailureofwordretrievalfrommemoryisthetip-of-the-tonguephenomenon.ThosewithAnomicaphasia(alsocalledNominalaphasiaorAnomia),however,doexperiencethetip-of-the-tonguephenomenononanongoingbasisduetodamagetothefrontalandparietallobesofthebrain. Memorydysfunctioncanalsooccurafterviralinfections.[104]ManypatientsrecoveringfromCOVID-19experiencememorylapses.Othervirusescanalsoelicitmemorydysfunction,includingSARS-CoV-1,MERS-CoV,Ebolavirusandeveninfluenzavirus.[104][105] Influencingfactors[edit] Interferencecanhampermemorizationandretrieval.Thereisretroactiveinterference,whenlearningnewinformationmakesithardertorecalloldinformation[106]andproactiveinterference,wherepriorlearningdisruptsrecallofnewinformation.Althoughinterferencecanleadtoforgetting,itisimportanttokeepinmindthattherearesituationswhenoldinformationcanfacilitatelearningofnewinformation.KnowingLatin,forinstance,canhelpanindividuallearnarelatedlanguagesuchasFrench–thisphenomenonisknownaspositivetransfer.[107] Stress[edit] Mainarticle:Effectsofstressonmemory Stresshasasignificanteffectonmemoryformationandlearning.Inresponsetostressfulsituations,thebrainreleaseshormonesandneurotransmitters(ex.glucocorticoidsandcatecholamines)whichaffectmemoryencodingprocessesinthehippocampus.Behaviouralresearchonanimalsshowsthatchronicstressproducesadrenalhormoneswhichimpactthehippocampalstructureinthebrainsofrats.[108]AnexperimentalstudybyGermancognitivepsychologistsL.SchwabeandO.Wolfdemonstrateshowlearningunderstressalsodecreasesmemoryrecallinhumans.[109]Inthisstudy,48healthyfemaleandmaleuniversitystudentsparticipatedineitherastresstestoracontrolgroup.Thoserandomlyassignedtothestresstestgrouphadahandimmersedinicecoldwater(thereputableSECPTor'SociallyEvaluatedColdPressorTest')foruptothreeminutes,whilebeingmonitoredandvideotaped.Boththestressandcontrolgroupswerethenpresentedwith32wordstomemorize.Twenty-fourhourslater,bothgroupsweretestedtoseehowmanywordstheycouldremember(freerecall)aswellashowmanytheycouldrecognizefromalargerlistofwords(recognitionperformance).Theresultsshowedaclearimpairmentofmemoryperformanceinthestresstestgroup,whorecalled30%fewerwordsthanthecontrolgroup.Theresearcherssuggestthatstressexperiencedduringlearningdistractspeoplebydivertingtheirattentionduringthememoryencodingprocess. However,memoryperformancecanbeenhancedwhenmaterialislinkedtothelearningcontext,evenwhenlearningoccursunderstress.AseparatestudybycognitivepsychologistsSchwabeandWolfshowsthatwhenretentiontestingisdoneinacontextsimilartoorcongruentwiththeoriginallearningtask(i.e.,inthesameroom),memoryimpairmentandthedetrimentaleffectsofstressonlearningcanbeattenuated.[110]Seventy-twohealthyfemaleandmaleuniversitystudents,randomlyassignedtotheSECPTstresstestortoacontrolgroup,wereaskedtorememberthelocationsof15pairsofpicturecards–acomputerizedversionofthecardgame"Concentration"or"Memory".Theroominwhichtheexperimenttookplacewasinfusedwiththescentofvanilla,asodourisastrongcueformemory.Retentiontestingtookplacethefollowingday,eitherinthesameroomwiththevanillascentagainpresent,orinadifferentroomwithoutthefragrance.Thememoryperformanceofsubjectswhoexperiencedstressduringtheobject-locationtaskdecreasedsignificantlywhentheyweretestedinanunfamiliarroomwithoutthevanillascent(anincongruentcontext);however,thememoryperformanceofstressedsubjectsshowednoimpairmentwhentheyweretestedintheoriginalroomwiththevanillascent(acongruentcontext).Allparticipantsintheexperiment,bothstressedandunstressed,performedfasterwhenthelearningandretrievalcontextsweresimilar.[111] Thisresearchontheeffectsofstressonmemorymayhavepracticalimplicationsforeducation,foreyewitnesstestimonyandforpsychotherapy:studentsmayperformbetterwhentestedintheirregularclassroomratherthananexamroom,eyewitnessesmayrecalldetailsbetteratthesceneofaneventthaninacourtroom,andpersonswithpost-traumaticstressmayimprovewhenhelpedtosituatetheirmemoriesofatraumaticeventinanappropriatecontext. Stressfullifeexperiencesmaybeacauseofmemorylossasapersonages.Glucocorticoidsthatarereleasedduringstresscausedamagetoneuronsthatarelocatedinthehippocampalregionofthebrain.Therefore,themorestressfulsituationsthatsomeoneencounters,themoresusceptibletheyaretomemorylosslateron.TheCA1neuronsfoundinthehippocampusaredestroyedduetoglucocorticoidsdecreasingthereleaseofglucoseandthereuptakeofglutamate.ThishighlevelofextracellularglutamateallowscalciumtoenterNMDAreceptorswhichinreturnkillsneurons.Stressfullifeexperiencescanalsocauserepressionofmemorieswhereapersonmovesanunbearablememorytotheunconsciousmind.[67]Thisdirectlyrelatestotraumaticeventsinone'spastsuchaskidnappings,beingprisonersofwarorsexualabuseasachild. Themorelongtermtheexposuretostressis,themoreimpactitmayhave.However,shorttermexposuretostressalsocausesimpairmentinmemorybyinterferingwiththefunctionofthehippocampus.Researchshowsthatsubjectsplacedinastressfulsituationforashortamountoftimestillhavebloodglucocorticoidlevelsthathaveincreaseddrasticallywhenmeasuredaftertheexposureiscompleted.Whensubjectsareaskedtocompletealearningtaskaftershorttermexposuretheyoftenhavedifficulties.Prenatalstressalsohinderstheabilitytolearnandmemorizebydisruptingthedevelopmentofthehippocampusandcanleadtounestablishedlongtermpotentiationintheoffspringofseverelystressedparents.Althoughthestressisappliedprenatally,theoffspringshowincreasedlevelsofglucocorticoidswhentheyaresubjectedtostresslateroninlife.[112]Oneexplanationforwhychildrenfromlowersocioeconomicbackgroundstendtodisplaypoorermemoryperformancethantheirhigher-incomepeersistheeffectsofstressaccumulatedoverthecourseofthelifetime.[113]Theeffectsoflowincomeonthedevelopinghippocampusisalsothoughtbemediatedbychronicstressresponseswhichmayexplainwhychildrenfromlowerandhigher-incomebackgroundsdifferintermsofmemoryperformance.[113] Sleep[edit] Mainarticle:Sleepandmemory Makingmemoriesoccursthroughathree-stepprocess,whichcanbeenhancedbysleep.Thethreestepsareasfollows: Acquisitionwhichistheprocessofstorageandretrievalofnewinformationinmemory Consolidation Recall Sleepaffectsmemoryconsolidation.Duringsleep,theneuralconnectionsinthebrainarestrengthened.Thisenhancesthebrain'sabilitiestostabilizeandretainmemories.Therehavebeenseveralstudieswhichshowthatsleepimprovestheretentionofmemory,asmemoriesareenhancedthroughactiveconsolidation.Systemconsolidationtakesplaceduringslow-wavesleep(SWS).[114][medicalcitationneeded]Thisprocessimplicatesthatmemoriesarereactivatedduringsleep,butthattheprocessdoesn'tenhanceeverymemory.Italsoimplicatesthatqualitativechangesaremadetothememorieswhentheyaretransferredtolong-termstoreduringsleep.Duringsleep,thehippocampusreplaystheeventsofthedayfortheneocortex.Theneocortexthenreviewsandprocessesmemories,whichmovesthemintolong-termmemory.Whenonedoesnotgetenoughsleepitmakesitmoredifficulttolearnastheseneuralconnectionsarenotasstrong,resultinginalowerretentionrateofmemories.Sleepdeprivationmakesithardertofocus,resultingininefficientlearning.[114]Furthermore,somestudieshaveshownthatsleepdeprivationcanleadtofalsememoriesasthememoriesarenotproperlytransferredtolong-termmemory. Oneoftheprimaryfunctionsofsleepisthoughttobetheimprovementoftheconsolidationofinformation,asseveralstudieshavedemonstratedthatmemorydependsongettingsufficientsleepbetweentrainingandtest.[115]Additionally,dataobtainedfromneuroimagingstudieshaveshownactivationpatternsinthesleepingbrainthatmirrorthoserecordedduringthelearningoftasksfromthepreviousday,[115]suggestingthatnewmemoriesmaybesolidifiedthroughsuchrehearsal.[116] Constructionforgeneralmanipulation[edit] Althoughpeopleoftenthinkthatmemoryoperateslikerecordingequipment,thisisnotthecase.Themolecularmechanismsunderlyingtheinductionandmaintenanceofmemoryareverydynamicandcomprisedistinctphasescoveringatimewindowfromsecondstoevenalifetime.[117]Infact,researchhasrevealedthatourmemoriesareconstructed:"currenthypothesessuggestthatconstructiveprocessesallowindividualstosimulateandimaginefutureepisodes,[118]happenings,andscenarios.Sincethefutureisnotanexactrepetitionofthepast,simulationoffutureepisodesrequiresacomplexsystemthatcandrawonthepastinamannerthatflexiblyextractsandrecombineselementsofpreviousexperiences–aconstructiveratherthanareproductivesystem."[72]Peoplecanconstructtheirmemorieswhentheyencodethemand/orwhentheyrecallthem.Toillustrate,consideraclassicstudyconductedbyElizabethLoftusandJohnPalmer(1974)[119]inwhichpeoplewereinstructedtowatchafilmofatrafficaccidentandthenaskedaboutwhattheysaw.Theresearchersfoundthatthepeoplewhowereasked,"Howfastwerethecarsgoingwhentheysmashedintoeachother?"gavehigherestimatesthanthosewhowereasked,"Howfastwerethecarsgoingwhentheyhiteachother?"Furthermore,whenaskedaweeklaterwhethertheyhadseenbrokenglassinthefilm,thosewhohadbeenaskedthequestionwithsmashedweretwicemorelikelytoreportthattheyhadseenbrokenglassthanthosewhohadbeenaskedthequestionwithhit(therewasnobrokenglassdepictedinthefilm).Thus,thewordingofthequestionsdistortedviewers'memoriesoftheevent.Importantly,thewordingofthequestionledpeopletoconstructdifferentmemoriesoftheevent–thosewhowereaskedthequestionwithsmashedrecalledamoreseriouscaraccidentthantheyhadactuallyseen.Thefindingsofthisexperimentwerereplicatedaroundtheworld,andresearchersconsistentlydemonstratedthatwhenpeoplewereprovidedwithmisleadinginformationtheytendedtomisremember,aphenomenonknownasthemisinformationeffect.[120] Researchhasrevealedthataskingindividualstorepeatedlyimagineactionsthattheyhaveneverperformedoreventsthattheyhaveneverexperiencedcouldresultinfalsememories.Forinstance,GoffandRoediger[121](1998)askedparticipantstoimaginethattheyperformedanact(e.g.,breakatoothpick)andthenlateraskedthemwhethertheyhaddonesuchathing.Findingsrevealedthatthoseparticipantswhorepeatedlyimaginedperformingsuchanactweremorelikelytothinkthattheyhadactuallyperformedthatactduringthefirstsessionoftheexperiment.Similarly,Garryandhercolleagues(1996)[122]askedcollegestudentstoreporthowcertaintheywerethattheyexperiencedanumberofeventsaschildren(e.g.,brokeawindowwiththeirhand)andthentwoweekslateraskedthemtoimaginefourofthoseevents.Theresearchersfoundthatone-fourthofthestudentsaskedtoimaginethefoureventsreportedthattheyhadactuallyexperiencedsucheventsaschildren.Thatis,whenaskedtoimaginetheeventstheyweremoreconfidentthattheyexperiencedtheevents. Researchreportedin2013revealedthatitispossibletoartificiallystimulatepriormemoriesandartificiallyimplantfalsememoriesinmice.Usingoptogenetics,ateamofRIKEN-MITscientistscausedthemicetoincorrectlyassociateabenignenvironmentwithapriorunpleasantexperiencefromdifferentsurroundings.Somescientistsbelievethatthestudymayhaveimplicationsinstudyingfalsememoryformationinhumans,andintreatingPTSDandschizophrenia.[123][124][medicalcitationneeded] Memoryreconsolidationiswhenpreviouslyconsolidatedmemoriesarerecalledorretrievedfromlong-termmemorytoyouractiveconsciousness.Duringthisprocess,memoriescanbefurtherstrengthenedandaddedtobutthereisalsoriskofmanipulationinvolved.Weliketothinkofourmemoriesassomethingstableandconstantwhentheyarestoredinlong-termmemorybutthisisn'tthecase.Therearealargenumberofstudiesthatfoundthatconsolidationofmemoriesisnotasingulareventbutareputthroughtheprocessagain,knownasreconsolidation.[125]Thisiswhenamemoryisrecalledorretrievedandplacedbackintoyourworkingmemory.Thememoryisnowopentomanipulationfromoutsidesourcesandthemisinformationeffectwhichcouldbeduetomisattributingthesourceoftheinconsistentinformation,withorwithoutanintactoriginalmemorytrace(LindsayandJohnson,1989).[126]Onethingthatcanbesureisthatmemoryismalleable. Thisnewresearchintotheconceptofreconsolidationhasopenedthedoortomethodstohelpthosewithunpleasantmemoriesorthosethatstrugglewithmemories.Anexampleofthisisifyouhadatrulyfrighteningexperienceandrecallthatmemoryinalessarousingenvironment,thememorywillbeweakenthenexttimeitisretrieved.[125]"Somestudiessuggestthatover-trainedorstronglyreinforcedmemoriesdonotundergoreconsolidationifreactivatedthefirstfewdaysaftertraining,butdobecomesensitivetoreconsolidationinterferencewithtime."[125]This,howeverdoesnotmeanthatallmemoryissusceptibletoreconsolidation.Thereisevidencetosuggestthatmemorythathasundergonestrongtrainingandwhetherornotisitintentionalislesslikelytoundergoreconsolidation.[127]Therewasfurthertestingdonewithratsandmazesthatshowedthatreactivatedmemoriesweremoresusceptibletomanipulation,inbothgoodandbadways,thannewlyformedmemories.[128]Itisstillnotknownwhetherornotthesearenewmemoriesformedandit'saninabilitytoretrievetheproperoneforthesituationorifit'sareconsolidatedmemory.Becausethestudyofreconsolidationisstillanewerconcept,thereisstilldebateonwhetheritshouldbeconsideredscientificallysound. Improving[edit] Mainarticle:Improvingmemory AUCLAresearchstudypublishedintheJune2008issueoftheAmericanJournalofGeriatricPsychiatryfoundthatpeoplecanimprovecognitivefunctionandbrainefficiencythroughsimplelifestylechangessuchasincorporatingmemoryexercises,healthyeating,physicalfitnessandstressreductionintotheirdailylives.Thisstudyexamined17subjects,(averageage53)withnormalmemoryperformance. Eightsubjectswereaskedtofollowa"brainhealthy"diet,relaxation,physical,andmentalexercise(brainteasersandverbalmemorytrainingtechniques).After14days,theyshowedgreaterwordfluency(notmemory)comparedtotheirbaselineperformance.Nolong-termfollow-upwasconducted;itisthereforeunclearifthisinterventionhaslastingeffectsonmemory.[129] Therearealooselyassociatedgroupofmnemonicprinciplesandtechniquesthatcanbeusedtovastlyimprovememoryknownastheartofmemory. TheInternationalLongevityCenterreleasedin2001areport[130]whichincludesinpages14–16recommendationsforkeepingthemindingoodfunctionalityuntiladvancedage.Someoftherecommendationsare: tostayintellectuallyactivethroughlearning trainingorreading tokeepphysicallyactivesotopromotebloodcirculationtothebrain tosocialize toreducestress tokeepsleeptimeregular toavoiddepressionoremotionalinstability toobservegoodnutrition. Memorizationisamethodoflearningthatallowsanindividualtorecallinformationverbatim.Rotelearningisthemethodmostoftenused.Methodsofmemorizingthingshavebeenthesubjectofmuchdiscussionovertheyearswithsomewriters,suchasCosmosRosselliususingvisualalphabets.Thespacingeffectshowsthatanindividualismorelikelytorememberalistofitemswhenrehearsalisspacedoveranextendedperiodoftime.Incontrasttothisiscramming:anintensivememorizationinashortperiodoftime.thespacingeffectisexploitedtoimprovememoryinspacedrepetitionflashcardtraining.AlsorelevantistheZeigarnikeffectwhichstatesthatpeoplerememberuncompletedorinterruptedtasksbetterthancompletedones.Theso-calledMethodoflociusesspatialmemorytomemorizenon-spatialinformation.[131] Inplants[edit] Plantslackaspecializedorgandevotedtomemoryretention,soplantmemoryhasbeenacontroversialtopicinrecentyears.Newadvancesinthefieldhaveidentifiedthepresenceofneurotransmittersinplants,addingtothehypothesisthatplantsarecapableofremembering.[132]Actionpotentials,aphysiologicalresponsecharacteristicofneurons,havebeenshowntohaveaninfluenceonplantsaswell,includinginwoundresponsesandphotosynthesis.[132]Inadditiontothesehomologousfeaturesofmemorysystemsinbothplantsandanimals,plantshavealsobeenobservedtoencode,storeandretrievebasicshort-termmemories. Oneofthemostwell-studiedplantstoshowrudimentarymemoryistheVenusflytrap.NativetothesubtropicalwetlandsoftheeasternUnitedStates,VenusFlyTrapshaveevolvedtheabilitytoobtainmeatforsustenance,likelyduetothelackofnitrogeninthesoil.[133]Thisisdonebytwotrap-formingleaftipsthatsnapshutoncetriggeredbyapotentialprey.Oneachlobe,threetriggershairsawaitstimulation.Inordertomaximizethebenefittocostratio,theplantenablesarudimentaryformofmemoryinwhichtwotriggerhairsmustbestimulatedwithin30secondsinordertoresultintrapclosure.[133]Thissystemensuresthatthetraponlycloseswhenpotentialpreyiswithingrasp. Thetimelapsebetweentriggerhairstimulationssuggeststhattheplantcanrememberaninitialstimuluslongenoughforasecondstimulustoinitiatetrapclosure.Thismemoryisn'tencodedinabrain,asplantslackthisspecializedorgan.Rather,informationisstoredintheformofcytoplasmiccalciumlevels.Thefirsttriggercausesasubthresholdcytoplasmiccalciuminflux.[133]Thisinitialtriggerisn'tenoughtoactivatetrapclosure,soasubsequentstimulusallowsforasecondaryinfluxofcalcium.Thelattercalciumrisesuperimposesontheinitialone,creatinganactionpotentialthatpassesthreshold,resultingintrapclosure.[133]Researchers,toprovethatanelectricalthresholdmustbemettostimulatetrapclosure,excitedasingletriggerhairwithaconstantmechanicalstimulususingAg/AgClelectrodes.[134]Thetrapclosedafteronlyafewseconds.Thisexperimentgaveevidencetodemonstratethattheelectricalthreshold,notnecessarilythenumberoftriggerhairstimulations,wasthecontributingfactorinVenusFlyTrapmemory.Ithasbeenshownthattrapclosurecanbeblockedusinguncouplersandinhibitorsofvoltage-gatedchannels.[134]Aftertrapclosure,theseelectricalsignalsstimulateglandularproductionofjasmonicacidandhydrolases,allowingfordigestionoftheprey.[135] Thefieldofplantneurobiologyhasgainedalargeamountofinterestoverthepastdecade,leadingtoaninfluxofresearchregardingplantmemory.AlthoughtheVenusflytrapisoneofthemorehighlystudied,manyotherplantsexhibitthecapacitytoremember,includingtheMimosapudicathroughanexperimentconductedbyMonicaGaglianoandcolleaguesin2013.[136]TostudytheMimosapudica,Gaglianodesignedanapparatuswithwhichpottedmimosaplantscouldberepeatedlydroppedthesamedistanceandatthesamespeed.Itwasobservedthattheplantsdefensiveresponseofcurlingupitsleavesdecreasedoverthe60timestheexperimentwasrepeatedperplant.Toconfirmthatthiswasamechanismofmemoryratherthanexhaustion,someoftheplantswereshakenpostexperimentanddisplayednormaldefensiveresponsesofleafcurling.Thisexperimentalsodemonstratedlong-termmemoryintheplants,asitwasrepeatedamonthlaterandtheplantswereobservedtoremainunfazedbythedropping.[136] Seealso[edit] Psychologyportal Medicineportal Adaptivememory,memorysystemsthathaveevolvedtohelpretainsurvival-and-fitnessinformation Animalmemory 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Memory-relatedresourcesfromtheNationalInstitutesofHealth MemoryatWikipedia'ssisterprojects DefinitionsfromWiktionaryMediafromCommonsNewsfromWikinewsQuotationsfromWikiquoteTextsfromWikisourceTextbooksfromWikibooksResourcesfromWikiversity Articlesrelatedtomemory vteHumanintelligencetopicsTypes Collective Emotional Intellectual Linguistic Multiple Social Spatial (visuospatial) Abilities,traits,andconstructs Cognition Communication Creativity Fluidandcrystallizedintelligence gfactor Intelligencequotient Knowledge Learning Memory Problemsolving Reasoning Thought (abstraction) Understanding Visualprocessing Modelsandtheories Cattell–Horn–Carrolltheory Fluidandcrystallizedintelligence Multiple-intelligencestheory Three-stratumtheory Triarchictheory PASStheory Areasofresearch Evolutionofhumanintelligence HeritabilityofIQ Psychometrics Intelligenceandenvironment /fertility /height /health /longevity /neuroscience /personality /race /sex Outlineofhumanintelligence /thought vteHumanmemoryBasicconcepts Encoding Storage Recall Attention Consolidation Neuroanatomy TypesSensory Echoic Eidetic Eyewitness Haptic Iconic Motorlearning Visual Short-term "TheMagicalNumberSeven,PlusorMinusTwo" Workingmemory Long-term Activerecall Autobiographical Explicit Declarative Episodic Semantic Flashbulb Hyperthymesia Implicit Meaningfullearning Personal-event Procedural Rotelearning Selectiveretention Tipofthetongue Forgetting Amnesia anterograde childhood post-traumatic psychogenic retrograde selective transientglobal Decaytheory Forgettingcurve Interferencetheory Memoryinhibition Motivatedforgetting Repressedmemory Retrieval-inducedforgetting Weaponfocus Memoryerrors Confabulation Cryptomnesia Hindsightbias Imaginationinflation Memorybiases Memoryconformity Misattributionofmemory Misinformationeffect Source-monitoringerror Falsememory Deese–Roediger–McDermottparadigm Falsememorysyndrome Memoryimplantation Lostinthemalltechnique Recovered-memorytherapy Researchmethods Exceptionalmemory Indirecttestsofmemory Memorydisorder Ingroups Collectivememory Politicsofmemory Culturalmemory Memoryandsocialinteractions Memoryconformity Transactivememory Othertopics Aging Artofmemory chunking mnemonic Effectsofalcohol Effectsofexercise Emotion Memoryimprovement Sleep Trauma Insociety Memorysport WorldMemoryChampionships ShasPollak Related Absent-mindedness Atkinson–Shiffrinmemorymodel Context-dependent&state-dependentmemory Childhoodmemory Exosomaticmemory Freerecall Intermediate-termmemory Involuntarymemory flashbacks LevelsofProcessingmodel Metamemory Musclememory Priming intertrial Prospective&retrospectivememory TheSevenSinsofMemory PeopleResearchers RichardC.Atkinson RobertA.Bjork StephenJ.Ceci SusanClancy HermannEbbinghaus SigmundFreud PatriciaGoldman-Rakic IvanIzquierdo MarciaK.Johnson EricKandel ElizabethLoftus GeoffreyLoftus JamesMcGaugh EleanorMaguire GeorgeArmitageMiller BrendaMilner LynnNadel HenryL.RoedigerIII DanielSchacter RichardShiffrin ArthurP.Shimamura LarrySquire SusumuTonegawa AnneTreisman EndelTulving RobertStickgold Patients HM KC CliveWearing Other JonathanHancock PaulR.McHugh DominicO'Brien BenPridmore CosmosRossellius AndriySlyusarchuk  Psychologyportal  Philosophyportal vtePhysiologyofthenervoussystemPrimarilyCNS Arousal Wakefulness Intracranialpressure Lateralizationofbrainfunction Sleep Memory PrimarilyPNS Reflex Sensation BothEvokedpotential Bereitschaftspotential P300 Auditoryevokedpotential Somatosensoryevokedpotentials Visualevokedpotential Othershortterm Neurotransmission Chronaxie Membranepotential Actionpotential Postsynapticpotential Excitatory Inhibitory Longterm Axoplasmictransport Neuroregeneration/Nerveregeneration Neuroplasticity/Synapticplasticity Long-termpotentiation Long-termdepression Other Myelinogenesis vteMentalprocessesCognition Awareness Criticalthinking Decision-making Comprehension Consciousness Imagination Intuition Problemsolving Perception Amodal Haptic(touch) Sound pitch harmonics speech Social Perceptionasinterpretation Visual Color RGBmodel Peripheral Depth Form Memory Encoding Storage Recall Consolidation Other Attention Highernervousactivity Intention Learning Mentalfatigue Mentalset Thinking Volition Authoritycontrol:Nationallibraries Spain France(data) Germany Israel UnitedStates Japan CzechRepublic Retrievedfrom"https://en.wikipedia.org/w/index.php?title=Memory&oldid=1109854307" 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