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ABObloodgroupsystemWikipedia,thefreeencyclopedia
ABObloodgroupsystem
FromWikipedia,thefreeencyclopedia
TheABObloodgroupsystemisthe
mostimportantbloodtypesystem(or
bloodgroupsystem)inhumanblood
transfusion.TheassociatedantiAand
antiBantibodiesareusuallyIgM
antibodies,whichareproducedinthe
firstyearsoflifebysensitizationto
environmentalsubstances,suchas
food,bacteria,andviruses.ABOblood
typesarealsopresentinsomeother
animals,forexamplerodentsandapes,
suchaschimpanzees,bonobos,and
gorillas.[1]
Contents
ABObloodgroupantigenspresentonredbloodcellsandIgM
antibodiespresentintheserum
1Historyofdiscoveriesofthe
bloodtypes
2Antigens
3RoleofABOantigensin
transfusionmedicine
3.1AlterationofABO
antigensfortransfusion
4Genetics
4.1Subgroups
4.2Distributionand
evolutionaryhistory
4.3Origintheories
5Normalroleinthebody
5.1Bleedingand
thrombosis(von
Willebrandfactor)
5.2Diseaserisks
5.3ABOhemolytic
diseaseofthenewborn
6Pseudoscience
7Seealso
8References
9Furtherreading
10Externallinks
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Historyofdiscoveriesofthebloodtypes
TheABObloodgroupsystemiswidelycreditedtohavebeendiscoveredby
theAustrianscientistKarlLandsteiner,whoidentifiedtheO,A,andB
bloodtypesin1900.[2]LandsteineroriginallydescribedtheObloodtypeas
type"C",andinpartsofEuropeitisrenderedas"0"(zero),signifyingthe
lackofAorBantigen.LandsteinerwasawardedtheNobelPrizein
PhysiologyorMedicinein1930forhiswork.AlfredvonDecastelloand
AdrianoSturlidiscoveredthefourthtype,AB,in1902.[3]
Duetoinadequatecommunicationatthetime,itwassubsequentlyfound
thattheCzechserologistJanJanskhadindependentlypioneeredthe
classificationofhumanbloodintofourgroups,[4]butLandsteiner's
independentdiscoveryhadbeenacceptedbythescientificworldwhile
Janskremainedtheninrelativeobscurity.However,in1921anAmerican
medicalcommissionacknowledgedJansk'sclassification.JanJanskis
nowadayscreditedwiththefirstclassificationofbloodintothefourtypes
(A,B,AB,0).
Jansk'sclassificationremainsinusetoday.InRussiaandstatesofthe
formerUSSRarebloodtypesO,A,B,andABarerespectively
designatedI,II,III,andIV.[5]ThedesignationAandBwith
referencetobloodgroupswasproposedbyLudwikHirszfeld.
InAmerica,W.L.Mosspublishedhisown(verysimilar)workin
1910.[6]
CzechserologistJanJansk
iscreditedwiththefirst
classificationofbloodinto
thefourtypes(A,B,AB,0)
Ukrainemarineuniformimprint,
showingthewearer'sbloodtypeas"B
(III)Rh+"
LudwikHirszfeldandE.vonDungerndiscoveredtheheritabilityof
ABObloodgroupsin191011.FelixBernsteindemonstratingthe
correctbloodgroupinheritancepatternofmultipleallelesatonelocusin1924.[7]WatkinsandMorgan,in
England,discoveredthattheABOepitopeswereconferredbysugars,tobespecific,Nacetylgalactosamine
fortheAtypeandgalactosefortheBtype.[8][9][10]AftermuchpublishedliteratureclaimingthattheABH
substanceswereallattachedtoglycosphingolipids,Finneetal.(1978)foundthatthehumanerythrocyte
glycoproteinscontainpolylactosaminechains[11]thatcontainsABHsubstancesattachedandrepresentthe
majorityoftheantigens.[12][13][14]ThemainglycoproteinscarryingtheABHantigenswereidentifiedtobe
theBand3andBand4.5proteinsandglycophorin.[15]Later,Yamamoto'sgroupshowedtheprecise
glycosyltransferasesetthatconferstheA,BandOepitopes.[16]
Antigens
ThecentralprincipleoftheABOsystemisthatantigensinthisinstance,sugarsphysicallyexposedonthe
exteriorofredbloodcellsdifferbetweenindividuals,whohaveimmunologicaltoleranceonlytoward
whatoccursintheirownbodies.Asaresult,manyhumansexpressisoantibodiesantibodiesagainst
isoantigens,naturalcomponentspresentinthebodiesofothermembersofthesamespeciesbutnot
themselves.IsoantibodiesmaybepresentagainsttheAand/orBantigensinpeoplewhodonotthemselves
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havethesameantigensintheirownblood.Theseantibodiesactashaemagglutinins,whichcauseblood
cellstoclumpandbreakapartiftheycarrytheforeignantigens.Thisharshresponse,thoughanadaptive
reactionusefulagainstinfection,cancausedeathwhenlargeamountsofsuchcellsareencounteredaftera
bloodtransfusion,acircumstancenotencounteredinnaturalselectionpriortomodernhistory.BecauseA
andBantigensarechemicallymodifiedfromaprecursorformthatisalsopresentintypeOindividuals,
peoplewithtypeAandBantigenscanacceptblood
fromtypeOindividuals.
AntiAandantiBantibodies(called
isohaemagglutinins),whicharenotpresentinthe
newborn,appearinthefirstyearsoflife.AntiAand
antiBantibodiesareusuallyIgMtype,whicharenot
abletopassthroughtheplacentatothefetalblood
circulation.OtypeindividualscanproduceIgGtype
ABOantibodies.
TheprecursortotheABObloodgroupantigens,
presentinpeopleofallcommonbloodtypes,iscalled
theHantigen.IndividualswiththerareBombay
Diagramshowingthecarbohydratechainsthat
phenotype(hh)donotexpressantigenHontheirred
determinetheABObloodgroup
bloodcells.AstheHantigenservesasaprecursorfor
producingAandBantigens,theabsenceoftheH
antigenmeansthattheindividualsalsolackAorBantigensaswell
(similartoObloodgroup).However,unlikeOgroup,theHantigenis
absent,hencetheindividualsproduceisoantibodiestoantigenHaswell
astobothAandBantigens.IftheyreceivebloodfromsomeonewithO
bloodgroup,theantiHantibodieswillbindtotheHantigenonthered
bloodcells('RBC')ofthedonorbloodanddestroytheRBCsby
complementmediatedlysis.Therefore,peoplewithBombayphenotype
Studentbloodtest.Threedrops
canreceivebloodonlyfromotherhhdonors(althoughtheycandonate
ofbloodaremixedwithantiB
asthoughtheyweretypeO).SomeindividualswiththebloodgroupA1
(left)andantiA(right)serum.
mayalsobeabletoproduceantiHantibodiesduetothecomplete
Agglutinationontherightside
conversionofalltheHantigentoA1antigen.
indicatesbloodtypeA.
ProductionoftheHantigen,oritsdeficiencyintheBombayphenotype,
iscontrolledattheHlocusonchromosome19.TheHlocusisnotthesamegeneastheABOlocus,butitis
epistatictotheABOlocus,providingthesubstratefortheAandBallelestomodify.[17]TheHlocus
containsthreeexonsthatspanmorethan5kbofgenomicDNA,andencodesthefucosyltransferasethat
producestheHantigenonRBCs.TheHantigenisacarbohydratesequencewithcarbohydrateslinked
mainlytoprotein(withaminorfractionattachedtoceramidemoiety).ItconsistsofachainofD
galactose,DNacetylglucosamine,Dgalactose,and2linked,Lfucose,thechainbeingattachedto
theproteinorceramide.
TheABOlocus,whichislocatedonchromosome9,containssevenexonsthatspanmorethan18kbof
genomicDNA.Exon7isthelargestandcontainsmostofthecodingsequence.TheABOlocushasthree
mainalleleicforms:A,B,andO.TheAalleleencodesaglycosyltransferasethatbondsN
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acetylgalactosaminetotheDgalactoseendoftheHantigen,producingtheAantigen.TheBalleleencodes
aglycosyltransferasethatbondsDgalactosetotheDgalactoseendoftheHantigen,creatingtheB
antigen.
InthecaseoftheOallele,whencomparedtotheAallele,exon6lacksonenucleotide(guanine),which
resultsinalossofenzymaticactivity.Thisdifference,whichoccursatposition261,causesaframeshift
thatresultsintheprematureterminationofthetranslationand,thus,degradationofthemRNA.Thisresults
intheHantigenremainingunchangedinthecaseofOgroups.
ThemajorityoftheABOantigensareexpressedontheendsoflongpolylactosaminechainsattached
mainlytoband3protein,theanionexchangeproteinoftheRBCmembrane,andaminorityoftheepitopes
areexpressedonneutralglycosphingolipid.
RoleofABOantigensintransfusionmedicine
ForablooddonorandrecipienttobeABOcompatibleforatransfusion,therecipientmustnotbeableto
produceAntiAorAntiBantibodiesthatcorrespondtotheAorBantigensonthesurfaceofthedonor's
redbloodcells(sincetheredbloodcellsareisolatedfromwholebloodbeforetransfusion,itisunimportant
whetherthedonorbloodhasantibodiesinitsplasma).Iftheantibodiesoftherecipient'sbloodandthe
antigensonthedonor'sredbloodcellsdocorrespond,thedonorbloodisrejected.Onrejection,the
recipientmayexperienceAcutehemolytictransfusionreaction(AHTR).
InadditiontotheABOsystem,theRhbloodgroupsystemcanaffecttransfusioncompatibility.An
individualiseitherpositiveornegativefortheRhfactorthisisdenotedbya'+'or''aftertheirABOtype.
BloodthatisRhnegativecanbetransfusedintoapersonwhoisRhpositive,butanRhnegativeindividual
cancreateantibodiesforRhpositiveRBCs.
Becauseofthis,theAB+bloodtypeisreferredtoasthe"universalrecipient",asitpossessesneitherAntiB
orAntiAantibodiesinitsplasma,andcanreceivebothRhpositiveandRhnegativeblood.Similarly,the
Obloodtypeiscalledthe"universaldonor"sinceitsredbloodcellshavenoAorBantigensandareRh
negative,nootherbloodtypewillrejectit.
IdentificationofABOandRhgenefrequenciesamonghumanpopulationshasvariousbenefitsin
transfusionmedicine,transplantationanddiseaserisk.[18]
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ABOandRhbloodtypedonationshowingmatchesbetweendonorandrecipienttypes
Donors
O+
Recipients
O+
A+
B+
AB+
A+
B+
AB+
AB
AB
AlterationofABOantigensfortransfusion
InApril2007,aninternationalteamofresearchersannouncedinthejournalNatureBiotechnologyan
inexpensiveandefficientwaytoconverttypesA,B,andABbloodintotypeO.[19]Thisisdonebyusing
glycosidaseenzymesfromspecificbacteriatostripthebloodgroupantigensfromredbloodcells.The
removalofAandBantigensstilldoesnotaddresstheproblemoftheRhesusbloodgroupantigenonthe
bloodcellsofRhesuspositiveindividuals,andsobloodfromRhesusnegativedonorsmustbeused.Patient
trialswillbeconductedbeforethemethodcanbereliedoninlivesituations.
Anotherapproachtothebloodantigenproblemisthemanufactureofartificialblood,whichcouldactasa
substituteinemergencies.[20]
Genetics
Bloodgroupsareinheritedfrombothparents.TheABObloodtypeiscontrolledbyasinglegene(theABO
gene)withthreetypesofallelesinferredfromclassicalgenetics:i,IA,andIB.Thegeneencodesa
glycosyltransferasethatis,anenzymethatmodifiesthecarbohydratecontentoftheredbloodcell
antigens.Thegeneislocatedonthelongarmoftheninthchromosome(9q34).
TheIAallelegivestypeA,IBgivestypeB,andigivestypeO.AsbothIAandIBaredominantoveri,only
iipeoplehavetypeOblood.IndividualswithIAIAorIAihavetypeAblood,andindividualswithIBIBorIBi
havetypeB.IAIBpeoplehavebothphenotypes,becauseAandBexpressaspecialdominancerelationship:
codominance,whichmeansthattypeAandBparentscanhaveanABchild.AcouplewithtypeAandtype
BcanalsohaveatypeOchildiftheyarebothheterozygous(IBi,IAi)ThecisABphenotypehasasingle
enzymethatcreatesbothAandBantigens.TheresultingredbloodcellsdonotusuallyexpressAorB
antigenatthesamelevelthatwouldbeexpectedoncommongroupA1orBredbloodcells,whichcanhelp
solvetheproblemofanapparentlygeneticallyimpossiblebloodgroup.[21]
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AandBarecodominant,givingtheAB
phenotype.
Bloodgroupinheritance
Bloodtype
AB
IAi(AO)
IAIA(AA)
IBi(BO)
IBIB(BB)
IAIB(AB)
OorA
OorB
AorB
OOOOOO
OO
AOOOAO
OO
AOAOAO
AO
BOOOBO
OO
BOBOBO
BO
AOBOAO
BO
IAi(AO)
OorA
OorA
BorAB
A,BorAB
AOAOOO
OO
AAAOAO
OO
AAAAAO
AO
O,A,Bor
AB
ABABBO
BO
AAABAO
BO
IAIA(AA)
AorAB
AB
AorAB
AOAOAO
AO
AAAOAA
AO
AAAAAA
AA
ABAOAB
AO
ABABAB
AB
AAABAA
AB
OorB
O,A,Bor
AB
ii(OO)
ABAOBO
OO
AorAB
OorB
A,BorAB
ABABAO
AO
BBBOBO
OO
BBBBBO
BO
ABBBAO
BO
IBi(BO)
BOBOOO
OO
IBIB(BB)
BorAB
AB
BorAB
BOBOBO
BO
ABBOAB
BO
ABABAB
AB
BBBOBB
BO
BBBBBB
BB
ABBBAB
BB
AorB
A,BorAB
AorAB
A,BorAB
BorAB
AOAOBO
BO
AAAOAB
BO
AAAAAB
AB
ABAOBB
BO
ABABBB
BB
A,B,or
AB
AB
ii(OO)
Genotype
O
IAIB(AB)
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ABBOAO
OO
AAABAB
BB
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Thetableabovesummarizesthevariousbloodgroupschildrenmayinheritfromtheirparents.[22][23]
Genotypesareshowninthesecondcolumnandinsmallprintfortheoffspring:AOandAAbothtestas
typeABOandBBtestastypeB.Thefourpossibilitiesrepresentthecombinationsobtainedwhenone
alleleistakenfromeachparenteachhasa25%chance,butsomeoccurmorethanonce.
Bloodgroupinheritancebyphenotypeonly
Bloodtype
AB
OorA
OorB
AorB
OorA
OorA
O,A,BorAB
A,BorAB
OorB
O,A,BorAB
OorB
A,BorAB
AB
AorB
A,BorAB
A,BorAB
A,BorAB
Historically,ABObloodtestswereusedinparentaltesting,butin1957only50%ofAmericanmenfalsely
accusedwereabletousethemasevidenceagainstpaternity.[24]Occasionally,thebloodtypesofchildren
arenotconsistentwithexpectationsforexample,atypeOchildcanbeborntoanABparentduetorare
situations,suchasBombayphenotypeandcisAB.[25]
Subgroups
TheAbloodtypecontainsabouttwentysubgroups,ofwhichA1andA2arethemostcommon(over99%).
A1makesupabout80%ofallAtypeblood,withA2makingupalmostalloftherest.[26]Thesetwo
subgroupsarenotalwaysinterchangeableasfarastransfusionisconcerned,assomeA2individuals
produceantibodiesagainsttheA1antigen.Complicationscansometimesariseinrarecaseswhentyping
theblood.[26]
WiththedevelopmentofDNAsequencing,ithasbeenpossibletoidentifyamuchlargernumberofalleles
attheABOlocus,eachofwhichcanbecategorizedasA,B,orOintermsofthereactiontotransfusion,but
whichcanbedistinguishedbyvariationsintheDNAsequence.Therearesixcommonallelesinwhite
individualsoftheABOgenethatproduceone'sbloodtype:[27][28]
A
Distributionandevolutionaryhistory
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ThedistributionofthebloodgroupsA,B,OandABvariesacrosstheworldaccordingtothepopulation.
Therearealsovariationsinbloodtypedistributionwithinhumansubpopulations.
IntheUK,thedistributionofbloodtypefrequenciesthroughthepopulationstillshowssomecorrelationto
thedistributionofplacenamesandtothesuccessiveinvasionsandmigrationsincludingNorsemens,Danes,
Saxons,Celts,andNormanswhocontributedthemorphemestotheplacenamesandthegenestothe
population.[30]
ThetwocommonOalleles,O01andO02,sharetheirfirst261nucleotideswiththegroupAalleleA01.[31]
However,unlikethegroupAallele,aguanosinebaseissubsequentlydeleted.Aprematurestopcodon
resultsfromthisframeshiftmutation.Thisvariantisfoundworldwide,andlikelypredateshuman
migrationfromAfrica.TheO01alleleisconsideredtopredatetheO02allele.
SomeevolutionarybiologiststheorizethattheIAalleleevolvedearliest,followedbyO(bythedeletionofa
singlenucleotide,shiftingthereadingframe)andthenIB.Thischronologyaccountsforthepercentageof
peopleworldwidewitheachbloodtype.Itisconsistentwiththeacceptedpatternsofearlypopulation
movementsandvaryingprevalentbloodtypesindifferentpartsoftheworld:forinstance,Bisvery
commoninpopulationsofAsiandescent,butrareinonesofWesternEuropeandescent.Anothertheory
statesthattherearefourmainlineagesoftheABOgeneandthatmutationscreatingtypeOhaveoccurred
atleastthreetimesinhumans.[32]Fromoldesttoyoungest,theselineagescomprisethefollowingalleles:
A101/A201/O09,B101,O02andO01.ThecontinuedpresenceoftheOallelesishypothesizedtobethe
resultofbalancingselection.[32]BoththeoriescontradictthepreviouslyheldtheorythattypeOblood
evolvedearliest.
Origintheories
Itispossiblethatfoodandenvironmentalantigens(bacterial,viral,orplantantigens)haveepitopessimilar
enoughtoAandBglycoproteinantigens.Theantibodiescreatedagainsttheseenvironmentalantigensin
thefirstyearsoflifecancrossreactwithABOincompatibleredbloodcellsthatitcomesincontactwith
duringbloodtransfusionlaterinlife.AntiAantibodiesarehypothesizedtooriginatefromimmune
responsetowardsinfluenzavirus,whoseepitopesaresimilarenoughtotheDNgalactosamineontheA
glycoproteintobeabletoelicitacrossreaction.AntiBantibodiesarehypothesizedtooriginatefrom
antibodiesproducedagainstGramnegativebacteria,suchasE.coli,crossreactingwiththeDgalactose
ontheBglycoprotein.[33]
HIVcanbeneutralizedininvitroexperimentsusingantibodiesagainstbloodgroupantigensspecifically
expressedontheHIVproducingcelllines.[34][35]
However,itismorelikelythattheforcedrivingevolutionofallelediversityissimplynegativefrequency
dependentselectioncellswithrarevariantsofmembraneantigensaremoreeasilydistinguishedbythe
immunesystemfrompathogenscarryingantigensfromotherhosts.Thus,individualspossessingraretypes
arebetterequippedtodetectpathogens.Thehighwithinpopulationdiversityobservedinhuman
populationswould,then,beaconsequenceofnaturalselectiononindividuals.[36]
Normalroleinthebody
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Thecarbohydratemoleculesonthesurfacesofredbloodcellshaverolesincellmembraneintegrity,cell
adhesion,membranetransportationofmolecules,andactingasreceptorsforextracellularligands,and
enzymes.ABOantigensarefoundhavingsimilarrolesonepithelialcellsaswellasredbloodcells.[37][38]
Bleedingandthrombosis(vonWillebrandfactor)
TheABOantigenisalsoexpressedonthevonWillebrandfactor(vWF)glycoprotein,[39]whichparticipates
inhemostasis(controlofbleeding).Infact,havingtypeObloodpredisposestobleeding,[40]as30%ofthe
totalgeneticvariationobservedinplasmavWFisexplainedbytheeffectoftheABObloodgroup,[41]and
individualswithgroupObloodnormallyhavesignificantlylowerplasmalevelsofvWF(andFactorVIII)
thandononOindividuals.[42][43]Inaddition,vWFisdegradedmorerapidlyduetothehigherprevalenceof
bloodgroupOwiththeCys1584variantofvWF(anaminoacidpolymorphisminVWF):[44]thegenefor
ADAMTS13(vWFcleavingprotease)mapstotheninthchromosome(9q34),thesamelocusasABOblood
type.HigherlevelsofvWFaremorecommonamongstpeoplewhohavehadischaemicstroke(fromblood
clotting)forthefirsttime.[45]Theresultsofthisstudyfoundthattheoccurrencewasnotaffectedby
ADAMTS13polymorphism,andtheonlysignificantgeneticfactorwastheperson'sbloodgroup.
Diseaserisks
ComparedtoOgroupindividuals,nonOgroup(A,AB,andB)individualshavea14%reducedriskof
squamouscellcarcinomaand4%reducedriskofbasalcellcarcinoma.[46]Conversely,typeObloodis
associatedwithareducedriskofpancreaticcancer.[47][48]TheBantigenlinkswithincreasedriskofovarian
cancer.[49]GastriccancerhasreportedtobemorecommoninbloodgroupAandleastingroupO.[50]
AccordingtoGlass,Holmgren,etal.,thoseintheObloodgrouphaveanincreasedriskofinfectionwith
cholera,andthoseOgroupindividualswhoareinfectedhavemoresevereinfections.Themechanisms
behindthisassociationwithcholeraarecurrentlyunclearintheliterature.[51]
ABOhemolyticdiseaseofthenewborn
ABObloodgroupincompatibilitiesbetweenthemotherandchilddoesnotusuallycausehemolyticdisease
ofthenewborn(HDN)becauseantibodiestotheABObloodgroupsareusuallyoftheIgMtype,whichdo
notcrosstheplacenta.However,inanOtypemother,IgGABOantibodiesareproducedandthebabycan
potentiallydevelopABOhemolyticdiseaseofthenewborn.
Pseudoscience
Duringthe1930s,connectingbloodgroupstopersonalitytypesbecamepopularinJapanandotherareasof
theworld.[52]Onthecontrary,therearesomepositivesciencestudies.[53][54]
Otherpopularbutunsupportedideasincludetheuseofabloodtypediet,claimsthatgroupAcausessevere
hangovers,groupOisassociatedwithperfectteeth,andthosewithbloodgroupA2havethehighestIQs.
Scientificevidenceinsupportoftheseconceptsisnonexistent.[55]
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Seealso
Artificialblood
Bloodtransfusion
Bombayphenotype
CisAB
Kiddbloodgroup
Rhbloodgroupsystem
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Furtherreading
DeanL(2005)."Chapter5:TheABObloodgroup."(http://www.ncbi.nlm.nih.gov/books/bv.fcgi?
rid=rbcantigen.chapter.ch05ABO).BloodGroupsandRedCellAntigens.Retrieved24March2007.
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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)at
BGMUTBloodGroupAntigenGeneMutationDatabaseatNCBI,NIH
ABObloodgroups,antibodiesandantigensexplained(https://www.youtube.com/watch?
v=u7DxZmLWDII)YouTubeeducationalvideo
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EncyclopaediaBritannica,ABObloodgroupsystem(http://www.britannica.com/eb/article
9003372/ABObloodgroupsystem)
NationalBloodTransfusionService(http://www.blood.co.uk/pages/world_blood.html)
MolecularGeneticBasisofABO(http://abobloodgroup.googlepages.com)
Retrievedfrom"http://en.wikipedia.org/w/index.php?title=ABO_blood_group_system&oldid=661578164"
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