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ABObloodgroupantigenspresentonredbloodcellsandIgMantibodiespresentintheserum
ABObloodgroupsystemFromWikipedia,thefreeencyclopedia
TheABObloodgroupsystemisthemostimportantbloodtypesystem(orbloodgroupsystem)inhumanbloodtransfusion.TheassociatedantiAandantiBantibodiesareusuallyIgMantibodies,whichareproducedinthefirstyearsoflifebysensitizationtoenvironmentalsubstances,suchasfood,bacteria,andviruses.ABObloodtypesarealsopresentinsomeotheranimals,forexamplerodentsandapes,suchaschimpanzees,bonobos,andgorillas.[1]
Contents
1Historyofdiscoveriesofthebloodtypes2Antigens3RoleofABOantigensintransfusionmedicine
3.1AlterationofABOantigensfortransfusion
4Genetics4.1Subgroups4.2Distributionandevolutionaryhistory4.3Origintheories
5Normalroleinthebody5.1Bleedingandthrombosis(vonWillebrandfactor)5.2Diseaserisks5.3ABOhemolyticdiseaseofthenewborn
6Pseudoscience7Seealso8References9Furtherreading10Externallinks
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CzechserologistJanJanskiscreditedwiththefirstclassificationofbloodintothefourtypes(A,B,AB,0)
Ukrainemarineuniformimprint,showingthewearer'sbloodtypeas"B(III)Rh+"
Historyofdiscoveriesofthebloodtypes
TheABObloodgroupsystemiswidelycreditedtohavebeendiscoveredbytheAustrianscientistKarlLandsteiner,whoidentifiedtheO,A,andBbloodtypesin1900.[2]LandsteineroriginallydescribedtheObloodtypeastype"C",andinpartsofEuropeitisrenderedas"0"(zero),signifyingthelackofAorBantigen.LandsteinerwasawardedtheNobelPrizeinPhysiologyorMedicinein1930forhiswork.AlfredvonDecastelloandAdrianoSturlidiscoveredthefourthtype,AB,in1902.[3]
Duetoinadequatecommunicationatthetime,itwassubsequentlyfoundthattheCzechserologistJanJanskhadindependentlypioneeredtheclassificationofhumanbloodintofourgroups,[4]butLandsteiner'sindependentdiscoveryhadbeenacceptedbythescientificworldwhileJanskremainedtheninrelativeobscurity.However,in1921anAmericanmedicalcommissionacknowledgedJansk'sclassification.JanJanskisnowadayscreditedwiththefirstclassificationofbloodintothefourtypes(A,B,AB,0).
Jansk'sclassificationremainsinusetoday.InRussiaandstatesoftheformerUSSRarebloodtypesO,A,B,andABarerespectivelydesignatedI,II,III,andIV.[5]ThedesignationAandBwithreferencetobloodgroupswasproposedbyLudwikHirszfeld.
InAmerica,W.L.Mosspublishedhisown(verysimilar)workin1910.[6]
LudwikHirszfeldandE.vonDungerndiscoveredtheheritabilityofABObloodgroupsin191011.FelixBernsteindemonstratingthecorrectbloodgroupinheritancepatternofmultipleallelesatonelocusin1924.[7]WatkinsandMorgan,inEngland,discoveredthattheABOepitopeswereconferredbysugars,tobespecific,NacetylgalactosaminefortheAtypeandgalactosefortheBtype.[8][9][10]AftermuchpublishedliteratureclaimingthattheABHsubstanceswereallattachedtoglycosphingolipids,Finneetal.(1978)foundthatthehumanerythrocyteglycoproteinscontainpolylactosaminechains[11]thatcontainsABHsubstancesattachedandrepresentthemajorityoftheantigens.[12][13][14]ThemainglycoproteinscarryingtheABHantigenswereidentifiedtobetheBand3andBand4.5proteinsandglycophorin.[15]Later,Yamamoto'sgroupshowedthepreciseglycosyltransferasesetthatconferstheA,BandOepitopes.[16]
Antigens
ThecentralprincipleoftheABOsystemisthatantigensinthisinstance,sugarsphysicallyexposedontheexteriorofredbloodcellsdifferbetweenindividuals,whohaveimmunologicaltoleranceonlytowardwhatoccursintheirownbodies.Asaresult,manyhumansexpressisoantibodiesantibodiesagainstisoantigens,naturalcomponentspresentinthebodiesofothermembersofthesamespeciesbutnotthemselves.IsoantibodiesmaybepresentagainsttheAand/orBantigensinpeoplewhodonotthemselves
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DiagramshowingthecarbohydratechainsthatdeterminetheABObloodgroup
Studentbloodtest.ThreedropsofbloodaremixedwithantiB(left)andantiA(right)serum.AgglutinationontherightsideindicatesbloodtypeA.
havethesameantigensintheirownblood.Theseantibodiesactashaemagglutinins,whichcausebloodcellstoclumpandbreakapartiftheycarrytheforeignantigens.Thisharshresponse,thoughanadaptivereactionusefulagainstinfection,cancausedeathwhenlargeamountsofsuchcellsareencounteredafterabloodtransfusion,acircumstancenotencounteredinnaturalselectionpriortomodernhistory.BecauseAandBantigensarechemicallymodifiedfromaprecursorformthatisalsopresentintypeOindividuals,peoplewithtypeAandBantigenscanacceptbloodfromtypeOindividuals.
AntiAandantiBantibodies(calledisohaemagglutinins),whicharenotpresentinthenewborn,appearinthefirstyearsoflife.AntiAandantiBantibodiesareusuallyIgMtype,whicharenotabletopassthroughtheplacentatothefetalbloodcirculation.OtypeindividualscanproduceIgGtypeABOantibodies.
TheprecursortotheABObloodgroupantigens,presentinpeopleofallcommonbloodtypes,iscalledtheHantigen.IndividualswiththerareBombayphenotype(hh)donotexpressantigenHontheirredbloodcells.AstheHantigenservesasaprecursorforproducingAandBantigens,theabsenceoftheHantigenmeansthattheindividualsalsolackAorBantigensaswell(similartoObloodgroup).However,unlikeOgroup,theHantigenisabsent,hencetheindividualsproduceisoantibodiestoantigenHaswellastobothAandBantigens.IftheyreceivebloodfromsomeonewithObloodgroup,theantiHantibodieswillbindtotheHantigenontheredbloodcells('RBC')ofthedonorbloodanddestroytheRBCsbycomplementmediatedlysis.Therefore,peoplewithBombayphenotypecanreceivebloodonlyfromotherhhdonors(althoughtheycandonateasthoughtheyweretypeO).SomeindividualswiththebloodgroupA1mayalsobeabletoproduceantiHantibodiesduetothecompleteconversionofalltheHantigentoA1antigen.
ProductionoftheHantigen,oritsdeficiencyintheBombayphenotype,iscontrolledattheHlocusonchromosome19.TheHlocusisnotthesamegeneastheABOlocus,butitisepistatictotheABOlocus,providingthesubstratefortheAandBallelestomodify.[17]TheHlocuscontainsthreeexonsthatspanmorethan5kbofgenomicDNA,andencodesthefucosyltransferasethatproducestheHantigenonRBCs.TheHantigenisacarbohydratesequencewithcarbohydrateslinkedmainlytoprotein(withaminorfractionattachedtoceramidemoiety).ItconsistsofachainofDgalactose,DNacetylglucosamine,Dgalactose,and2linked,Lfucose,thechainbeingattachedtotheproteinorceramide.
TheABOlocus,whichislocatedonchromosome9,containssevenexonsthatspanmorethan18kbofgenomicDNA.Exon7isthelargestandcontainsmostofthecodingsequence.TheABOlocushasthreemainalleleicforms:A,B,andO.TheAalleleencodesaglycosyltransferasethatbondsN
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acetylgalactosaminetotheDgalactoseendoftheHantigen,producingtheAantigen.TheBalleleencodesaglycosyltransferasethatbondsDgalactosetotheDgalactoseendoftheHantigen,creatingtheBantigen.
InthecaseoftheOallele,whencomparedtotheAallele,exon6lacksonenucleotide(guanine),whichresultsinalossofenzymaticactivity.Thisdifference,whichoccursatposition261,causesaframeshiftthatresultsintheprematureterminationofthetranslationand,thus,degradationofthemRNA.ThisresultsintheHantigenremainingunchangedinthecaseofOgroups.
ThemajorityoftheABOantigensareexpressedontheendsoflongpolylactosaminechainsattachedmainlytoband3protein,theanionexchangeproteinoftheRBCmembrane,andaminorityoftheepitopesareexpressedonneutralglycosphingolipid.
RoleofABOantigensintransfusionmedicine
ForablooddonorandrecipienttobeABOcompatibleforatransfusion,therecipientmustnotbeabletoproduceAntiAorAntiBantibodiesthatcorrespondtotheAorBantigensonthesurfaceofthedonor'sredbloodcells(sincetheredbloodcellsareisolatedfromwholebloodbeforetransfusion,itisunimportantwhetherthedonorbloodhasantibodiesinitsplasma).Iftheantibodiesoftherecipient'sbloodandtheantigensonthedonor'sredbloodcellsdocorrespond,thedonorbloodisrejected.Onrejection,therecipientmayexperienceAcutehemolytictransfusionreaction(AHTR).
InadditiontotheABOsystem,theRhbloodgroupsystemcanaffecttransfusioncompatibility.AnindividualiseitherpositiveornegativefortheRhfactorthisisdenotedbya'+'or''aftertheirABOtype.BloodthatisRhnegativecanbetransfusedintoapersonwhoisRhpositive,butanRhnegativeindividualcancreateantibodiesforRhpositiveRBCs.
Becauseofthis,theAB+bloodtypeisreferredtoasthe"universalrecipient",asitpossessesneitherAntiBorAntiAantibodiesinitsplasma,andcanreceivebothRhpositiveandRhnegativeblood.Similarly,theObloodtypeiscalledthe"universaldonor"sinceitsredbloodcellshavenoAorBantigensandareRhnegative,nootherbloodtypewillrejectit.
IdentificationofABOandRhgenefrequenciesamonghumanpopulationshasvariousbenefitsintransfusionmedicine,transplantationanddiseaserisk.[18]
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ABOandRhbloodtypedonationshowingmatchesbetweendonorandrecipienttypesDonors
O+ A+ B+ AB+ O A B AB
Recipients
O+ A+ B+ AB+ O A B AB
AlterationofABOantigensfortransfusion
InApril2007,aninternationalteamofresearchersannouncedinthejournalNatureBiotechnologyaninexpensiveandefficientwaytoconverttypesA,B,andABbloodintotypeO.[19]Thisisdonebyusingglycosidaseenzymesfromspecificbacteriatostripthebloodgroupantigensfromredbloodcells.TheremovalofAandBantigensstilldoesnotaddresstheproblemoftheRhesusbloodgroupantigenonthebloodcellsofRhesuspositiveindividuals,andsobloodfromRhesusnegativedonorsmustbeused.Patienttrialswillbeconductedbeforethemethodcanbereliedoninlivesituations.
Anotherapproachtothebloodantigenproblemisthemanufactureofartificialblood,whichcouldactasasubstituteinemergencies.[20]
Genetics
Bloodgroupsareinheritedfrombothparents.TheABObloodtypeiscontrolledbyasinglegene(theABOgene)withthreetypesofallelesinferredfromclassicalgenetics:i,IA,andIB.Thegeneencodesaglycosyltransferasethatis,anenzymethatmodifiesthecarbohydratecontentoftheredbloodcellantigens.Thegeneislocatedonthelongarmoftheninthchromosome(9q34).
TheIAallelegivestypeA,IBgivestypeB,andigivestypeO.AsbothIAandIBaredominantoveri,onlyiipeoplehavetypeOblood.IndividualswithIAIAorIAihavetypeAblood,andindividualswithIBIBorIBihavetypeB.IAIBpeoplehavebothphenotypes,becauseAandBexpressaspecialdominancerelationship:codominance,whichmeansthattypeAandBparentscanhaveanABchild.AcouplewithtypeAandtypeBcanalsohaveatypeOchildiftheyarebothheterozygous(IBi,IAi)ThecisABphenotypehasasingleenzymethatcreatesbothAandBantigens.TheresultingredbloodcellsdonotusuallyexpressAorBantigenatthesamelevelthatwouldbeexpectedoncommongroupA1orBredbloodcells,whichcanhelp
solvetheproblemofanapparentlygeneticallyimpossiblebloodgroup.[21]
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AandBarecodominant,givingtheABphenotype.
BloodgroupinheritanceBloodtype O A B AB
Genotype ii(OO) IAi(AO) IAIA(AA) IBi(BO) IBIB(BB) IAIB(AB)
O ii(OO)O
OOOOOOOO
OorAAOOOAO
OO
AAOAOAO
AO
OorBBOOOBO
OO
BBOBOBO
BO
AorBAOBOAO
BO
A
IAi(AO)OorA
AOAOOOOO
OorAAAAOAO
OO
AAAAAAO
AO
O,A,BorAB
ABAOBOOO
BorABABABBO
BO
A,BorABAAABAO
BO
IAIA(AA)A
AOAOAOAO
AAAAOAA
AO
AAAAAAA
AA
AorABABAOAB
AO
ABABABAB
AB
AorABAAABAA
AB
B
IBi(BO)OorB
BOBOOOOO
O,A,BorAB
ABBOAOOO
AorABABABAO
AO
OorBBBBOBO
OO
BBBBBBO
BO
A,BorABABBBAO
BO
IBIB(BB)B
BOBOBOBO
BorABABBOAB
BO
ABABABAB
AB
BBBBOBB
BO
BBBBBBB
BB
BorABABBBAB
BB
AB IAIB(AB)AorB
AOAOBOBO
A,BorABAAAOAB
BO
AorABAAAAAB
AB
A,BorABABAOBB
BO
BorABABABBB
BB
A,B,orAB
AAABABBB
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BloodgroupinheritancebyphenotypeonlyBloodtype O A B AB
O O OorA OorB AorBA OorA OorA O,A,BorAB A,BorABB OorB O,A,BorAB OorB A,BorABAB AorB A,BorAB A,BorAB A,BorAB
Thetableabovesummarizesthevariousbloodgroupschildrenmayinheritfromtheirparents.[22][23]Genotypesareshowninthesecondcolumnandinsmallprintfortheoffspring:AOandAAbothtestastypeABOandBBtestastypeB.Thefourpossibilitiesrepresentthecombinationsobtainedwhenonealleleistakenfromeachparenteachhasa25%chance,butsomeoccurmorethanonce.
Historically,ABObloodtestswereusedinparentaltesting,butin1957only50%ofAmericanmenfalselyaccusedwereabletousethemasevidenceagainstpaternity.[24]Occasionally,thebloodtypesofchildrenarenotconsistentwithexpectationsforexample,atypeOchildcanbeborntoanABparentduetoraresituations,suchasBombayphenotypeandcisAB.[25]
Subgroups
TheAbloodtypecontainsabouttwentysubgroups,ofwhichA1andA2arethemostcommon(over99%).A1makesupabout80%ofallAtypeblood,withA2makingupalmostalloftherest.[26]Thesetwosubgroupsarenotalwaysinterchangeableasfarastransfusionisconcerned,assomeA2individualsproduceantibodiesagainsttheA1antigen.Complicationscansometimesariseinrarecaseswhentypingtheblood.[26]
WiththedevelopmentofDNAsequencing,ithasbeenpossibletoidentifyamuchlargernumberofallelesattheABOlocus,eachofwhichcanbecategorizedasA,B,orOintermsofthereactiontotransfusion,butwhichcanbedistinguishedbyvariationsintheDNAsequence.TherearesixcommonallelesinwhiteindividualsoftheABOgenethatproduceone'sbloodtype:[27][28]
A B OA101(A1)A201(A2)
B101(B1) O01(O1)O02(O1v)O03(O2)
Thesamestudyalsoidentified18rarealleles,whichgenerallyhaveaweakerglycosylationactivity.PeoplewithweakallelesofAcansometimesexpressantiAantibodies,thoughtheseareusuallynotclinicallysignificantastheydonotstablyinteractwiththeantigenatbodytemperature.[29]
CisABisanotherrarevariant,inwhichAandBgenesaretransmittedtogetherfromasingleparent.
Distributionandevolutionaryhistory
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ThedistributionofthebloodgroupsA,B,OandABvariesacrosstheworldaccordingtothepopulation.Therearealsovariationsinbloodtypedistributionwithinhumansubpopulations.
IntheUK,thedistributionofbloodtypefrequenciesthroughthepopulationstillshowssomecorrelationtothedistributionofplacenamesandtothesuccessiveinvasionsandmigrationsincludingNorsemens,Danes,Saxons,Celts,andNormanswhocontributedthemorphemestotheplacenamesandthegenestothepopulation.[30]
ThetwocommonOalleles,O01andO02,sharetheirfirst261nucleotideswiththegroupAalleleA01.[31]However,unlikethegroupAallele,aguanosinebaseissubsequentlydeleted.Aprematurestopcodonresultsfromthisframeshiftmutation.Thisvariantisfoundworldwide,andlikelypredateshumanmigrationfromAfrica.TheO01alleleisconsideredtopredatetheO02allele.
SomeevolutionarybiologiststheorizethattheIAalleleevolvedearliest,followedbyO(bythedeletionofasinglenucleotide,shiftingthereadingframe)andthenIB.Thischronologyaccountsforthepercentageofpeopleworldwidewitheachbloodtype.Itisconsistentwiththeacceptedpatternsofearlypopulationmovementsandvaryingprevalentbloodtypesindifferentpartsoftheworld:forinstance,BisverycommoninpopulationsofAsiandescent,butrareinonesofWesternEuropeandescent.AnothertheorystatesthattherearefourmainlineagesoftheABOgeneandthatmutationscreatingtypeOhaveoccurredatleastthreetimesinhumans.[32]Fromoldesttoyoungest,theselineagescomprisethefollowingalleles:A101/A201/O09,B101,O02andO01.ThecontinuedpresenceoftheOallelesishypothesizedtobetheresultofbalancingselection.[32]BoththeoriescontradictthepreviouslyheldtheorythattypeObloodevolvedearliest.
Origintheories
Itispossiblethatfoodandenvironmentalantigens(bacterial,viral,orplantantigens)haveepitopessimilarenoughtoAandBglycoproteinantigens.TheantibodiescreatedagainsttheseenvironmentalantigensinthefirstyearsoflifecancrossreactwithABOincompatibleredbloodcellsthatitcomesincontactwithduringbloodtransfusionlaterinlife.AntiAantibodiesarehypothesizedtooriginatefromimmuneresponsetowardsinfluenzavirus,whoseepitopesaresimilarenoughtotheDNgalactosamineontheAglycoproteintobeabletoelicitacrossreaction.AntiBantibodiesarehypothesizedtooriginatefromantibodiesproducedagainstGramnegativebacteria,suchasE.coli,crossreactingwiththeDgalactoseontheBglycoprotein.[33]
HIVcanbeneutralizedininvitroexperimentsusingantibodiesagainstbloodgroupantigensspecificallyexpressedontheHIVproducingcelllines.[34][35]
However,itismorelikelythattheforcedrivingevolutionofallelediversityissimplynegativefrequencydependentselectioncellswithrarevariantsofmembraneantigensaremoreeasilydistinguishedbytheimmunesystemfrompathogenscarryingantigensfromotherhosts.Thus,individualspossessingraretypesarebetterequippedtodetectpathogens.Thehighwithinpopulationdiversityobservedinhumanpopulationswould,then,beaconsequenceofnaturalselectiononindividuals.[36]
Normalroleinthebody
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Thecarbohydratemoleculesonthesurfacesofredbloodcellshaverolesincellmembraneintegrity,celladhesion,membranetransportationofmolecules,andactingasreceptorsforextracellularligands,andenzymes.ABOantigensarefoundhavingsimilarrolesonepithelialcellsaswellasredbloodcells.[37][38]
Bleedingandthrombosis(vonWillebrandfactor)
TheABOantigenisalsoexpressedonthevonWillebrandfactor(vWF)glycoprotein,[39]whichparticipatesinhemostasis(controlofbleeding).Infact,havingtypeObloodpredisposestobleeding,[40]as30%ofthetotalgeneticvariationobservedinplasmavWFisexplainedbytheeffectoftheABObloodgroup,[41]andindividualswithgroupObloodnormallyhavesignificantlylowerplasmalevelsofvWF(andFactorVIII)thandononOindividuals.[42][43]Inaddition,vWFisdegradedmorerapidlyduetothehigherprevalenceofbloodgroupOwiththeCys1584variantofvWF(anaminoacidpolymorphisminVWF):[44]thegeneforADAMTS13(vWFcleavingprotease)mapstotheninthchromosome(9q34),thesamelocusasABObloodtype.HigherlevelsofvWFaremorecommonamongstpeoplewhohavehadischaemicstroke(frombloodclotting)forthefirsttime.[45]TheresultsofthisstudyfoundthattheoccurrencewasnotaffectedbyADAMTS13polymorphism,andtheonlysignificantgeneticfactorwastheperson'sbloodgroup.
Diseaserisks
ComparedtoOgroupindividuals,nonOgroup(A,AB,andB)individualshavea14%reducedriskofsquamouscellcarcinomaand4%reducedriskofbasalcellcarcinoma.[46]Conversely,typeObloodisassociatedwithareducedriskofpancreaticcancer.[47][48]TheBantigenlinkswithincreasedriskofovariancancer.[49]GastriccancerhasreportedtobemorecommoninbloodgroupAandleastingroupO.[50]
AccordingtoGlass,Holmgren,etal.,thoseintheObloodgrouphaveanincreasedriskofinfectionwithcholera,andthoseOgroupindividualswhoareinfectedhavemoresevereinfections.Themechanismsbehindthisassociationwithcholeraarecurrentlyunclearintheliterature.[51]
ABOhemolyticdiseaseofthenewborn
ABObloodgroupincompatibilitiesbetweenthemotherandchilddoesnotusuallycausehemolyticdiseaseofthenewborn(HDN)becauseantibodiestotheABObloodgroupsareusuallyoftheIgMtype,whichdonotcrosstheplacenta.However,inanOtypemother,IgGABOantibodiesareproducedandthebabycanpotentiallydevelopABOhemolyticdiseaseofthenewborn.
Pseudoscience
Duringthe1930s,connectingbloodgroupstopersonalitytypesbecamepopularinJapanandotherareasoftheworld.[52]Onthecontrary,therearesomepositivesciencestudies.[53][54]
Otherpopularbutunsupportedideasincludetheuseofabloodtypediet,claimsthatgroupAcausesseverehangovers,groupOisassociatedwithperfectteeth,andthosewithbloodgroupA2havethehighestIQs.Scientificevidenceinsupportoftheseconceptsisnonexistent.[55]
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Seealso
ArtificialbloodBloodtransfusionBombayphenotypeCisABKiddbloodgroupRhbloodgroupsystem
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Furtherreading
DeanL(2005)."Chapter5:TheABObloodgroup."(http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=rbcantigen.chapter.ch05ABO).BloodGroupsandRedCellAntigens.Retrieved24March2007.FarrA(1April1979)."Bloodgroupserologythefirstfourdecades(19001939)"(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1082436).MedHist23(2):21526.doi:10.1017/s0025727300051383(https://dx.doi.org/10.1017%2Fs0025727300051383).PMC1082436(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1082436).PMID381816(https://www.ncbi.nlm.nih.gov/pubmed/381816).
Externallinks
ABO(http://www.ncbi.nlm.nih.gov/gv/mhc/xslcgi.cgi?cmd=bgmut/systems_info&system=abo)atBGMUTBloodGroupAntigenGeneMutationDatabaseatNCBI,NIHABObloodgroups,antibodiesandantigensexplained(https://www.youtube.com/watch?v=u7DxZmLWDII)YouTubeeducationalvideo
susceptibilitytopancreaticcancer"(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2839871).NatureGenetics41(9):986990.doi:10.1038/ng.429(https://dx.doi.org/10.1038%2Fng.429).PMC2839871(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2839871).PMID19648918(https://www.ncbi.nlm.nih.gov/pubmed/19648918).
49. Gates,MAWolpin,BMCramer,DWHankinson,SETworoger,SS(2010)."ABObloodgroupandincidenceofepithelialovariancancer"(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2946962).InternationalJournalofCancer.JournalInternationalDuCancer128(2):482486.doi:10.1002/ijc.25339(https://dx.doi.org/10.1002%2Fijc.25339).PMC2946962(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2946962).PMID20309936(https://www.ncbi.nlm.nih.gov/pubmed/20309936).
50. Aird,IBentall,HHRoberts,JA(1953)."ArelationshipbetweencancerofstomachandtheABObloodgroups"(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2015995).BritishMedicalJournal1(4814):799801.doi:10.1136/bmj.1.4814.799(https://dx.doi.org/10.1136%2Fbmj.1.4814.799).PMC2015995(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2015995).PMID13032504(https://www.ncbi.nlm.nih.gov/pubmed/13032504).
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52. AmericanRedCross,SouthernCaliforniaBloodServicesRegion(n.d.)."AnswerstoCommonlyAskedQuestionsAboutBloodandBloodBanking"(http://www.socalredcross.org/pdf/BloodThe.pdf)(PDF).Blood:theBasics:4.Archived(http://web.archive.org/web/20071129152741/http://www.socalredcross.org/pdf/BloodThe.pdf)(PDF)fromtheoriginalon29November2007.Retrieved16November2007.
53. SungIlRyu,YoungWooSohn(2007),AReviewofSociocultural,Behavioral,BiochemicalAnalysesonABOBloodGroupsTypology,TheKoreanJournalofSocialandPersonalityPsychology.
54. DonnaK.Hobgood(2011),PersonalitytraitsofaggressionsubmissivenessandperfectionismassociatewithABObloodgroupsthroughcatecholamineactivities(http://www.sciencedirect.com/science/article/pii/S0306987711002106),MedicalHypotheses,77(2):294300.
55. Klein,HarveyG(7March2005)."WhyDoPeopleHaveDifferentBloodTypes?"(http://www.sciam.com/article.cfm?id=whydopeoplehavediffer).ScientificAmerican.Retrieved16November2007.
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EncyclopaediaBritannica,ABObloodgroupsystem(http://www.britannica.com/eb/article9003372/ABObloodgroupsystem)NationalBloodTransfusionService(http://www.blood.co.uk/pages/world_blood.html)MolecularGeneticBasisofABO(http://abobloodgroup.googlepages.com)
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