You are on page 1of 14

5/12/2015

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

http://en.wikipedia.org/wiki/ABO_blood_group_system

1/14

5/12/2015

ABObloodgroupsystemWikipedia,thefreeencyclopedia

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
http://en.wikipedia.org/wiki/ABO_blood_group_system

2/14

5/12/2015

ABObloodgroupsystemWikipedia,thefreeencyclopedia

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

http://en.wikipedia.org/wiki/ABO_blood_group_system

3/14

5/12/2015

ABObloodgroupsystemWikipedia,thefreeencyclopedia

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]

http://en.wikipedia.org/wiki/ABO_blood_group_system

4/14

5/12/2015

ABObloodgroupsystemWikipedia,thefreeencyclopedia

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]

http://en.wikipedia.org/wiki/ABO_blood_group_system

5/14

5/12/2015

ABObloodgroupsystemWikipedia,thefreeencyclopedia

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)

http://en.wikipedia.org/wiki/ABO_blood_group_system

ABBOAO
OO

AAABAB
BB

6/14

5/12/2015

ABObloodgroupsystemWikipedia,thefreeencyclopedia

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

A101(A1) B101(B1) O01(O1)


A201(A2)
O02(O1v)
O03(O2)
Thesamestudyalsoidentified18rarealleles,whichgenerallyhaveaweakerglycosylationactivity.People
withweakallelesofAcansometimesexpressantiAantibodies,thoughtheseareusuallynotclinically
significantastheydonotstablyinteractwiththeantigenatbodytemperature.[29]
CisABisanotherrarevariant,inwhichAandBgenesaretransmittedtogetherfromasingleparent.

Distributionandevolutionaryhistory
http://en.wikipedia.org/wiki/ABO_blood_group_system

7/14

5/12/2015

ABObloodgroupsystemWikipedia,thefreeencyclopedia

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
http://en.wikipedia.org/wiki/ABO_blood_group_system

8/14

5/12/2015

ABObloodgroupsystemWikipedia,thefreeencyclopedia

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]

http://en.wikipedia.org/wiki/ABO_blood_group_system

9/14

5/12/2015

ABObloodgroupsystemWikipedia,thefreeencyclopedia

Seealso
Artificialblood
Bloodtransfusion
Bombayphenotype
CisAB
Kiddbloodgroup
Rhbloodgroupsystem

References
1. Maton,AntheaJeanHopkins,CharlesWilliamMcLaughlin,SusanJohnson,MaryannaQuonWarner,David
LaHart,JillD.Wright(1993).HumanBiologyandHealth.EnglewoodCliffs,NewJersey,USA:PrenticeHall.
ISBN0139811761.
2. LandsteinerK(1900)."ZurKenntnisderantifermentativen,lytischenundagglutinierendenWirkungendes
BlutserumsundderLymphe".ZentralblattBakteriologie27:35762.
3. vonDecastelloA,SturliA(1902)."UeberdieIsoagglutinineimSerumgesunderundkrankerMenschen".Mfinch
medWschr49:10905.
4. JanskJ(1907)."Haematologickstudieu.psychotiku".Sborn.Klinick(inCzech)8:85139.
5. ErbIH(1May1940)."BloodGroupClassifications,aPleaforUniformity"
(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC537907).CanadianMedicalAssociationJournal42(5):418
21.PMC537907(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC537907).PMID20321693
(https://www.ncbi.nlm.nih.gov/pubmed/20321693).
6. MossWL(1910)."Studiesonisoagglutininsandisohemolysins".BulletinJohnsHopkinsHospital21:6370.
7. CrowJ(1993)."FelixBernsteinandthefirsthumanmarkerlocus"
(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1205297).Genetics133(1):47.PMC1205297
(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1205297).PMID8417988
(https://www.ncbi.nlm.nih.gov/pubmed/8417988).
8. Morgan,W.T.J.&Watkins,W.M.Br.Med.Bull.25,3034(1969)
9. Watkins,W.M.AdvancesinHumanGeneticsVol.10(edsHarris,H.&Hirschhorn,K.)1136(Plenum,New
York,1980)
10. Watkins,W.M.&Morgan,W.T.J.VoxSang.4,97119(1959).
11. Finne,Krusius,Rauvala,Kekomki,Myllyl,FEBSleft.89:111115(1978)
12. Krusius,Finne,Rauvala,Eur.J.Biochem.92:289300(1978)
13. Jrnefelt,Rush,Li,Laine,J.Biol.Chem.253:80068009(1978)
14. LaineandRushinMolecularImmunologyofComplexCarbohydrates(A.Wu,E.Kabat,Eds.)Plenum
PublishingCorporation,N.Y.NY(1988)
15. Finne,Eur.J.Biochem.104:181189(1980)
16. Yamamoto,etal.,MoleculargeneticbasisofthehistobloodgroupABOsystem,Nature345:229233(1990)
17. BenjaminA.Pierce(20080215).TransmissionandPopulationGenetics(http://books.google.com/books?
id=UzYiLVvaNwYC&pg=PA108).ISBN9781429211185.
18. Fareed,MohdHussain,RuqaiyaShah,AhsanaAfzal,Mohammad(2014)."A1A2BOandRhgenefrequencies
amongsixpopulationsofJammuandKashmir,India".TransfusionandApheresisScience50(2):247.
doi:10.1016/j.transci.2014.01.014(https://dx.doi.org/10.1016%2Fj.transci.2014.01.014).PMID24485956
(https://www.ncbi.nlm.nih.gov/pubmed/24485956).
19. Liu,QPSulzenbacherG,YuanH,BennettEP,PietzG,SaundersK,SpenceJ,NudelmanE,LeverySB,White
T,NeveuJM,LaneWS,BourneY,OlssonML,HenrissatB,ClausenH(April2007)."Bacterialglycosidases
fortheproductionofuniversalredbloodcells"(http://www.nature.com/nbt/journal/v25/n4/abs/nbt1298.html).
NatBiotechnol25(4):45464.doi:10.1038/nbt1298(https://dx.doi.org/10.1038%2Fnbt1298).PMID17401360
(https://www.ncbi.nlm.nih.gov/pubmed/17401360).
20. BBC(http://news.bbc.co.uk/2/hi/uk_news/england/north_yorkshire/6645923.stm)
21. YazerM,OlssonM,PalcicMOlssonPalcic(2006)."ThecisABbloodgroupphenotype:fundamentallessons
http://en.wikipedia.org/wiki/ABO_blood_group_system
10/14

5/12/2015

ABObloodgroupsystemWikipedia,thefreeencyclopedia

21. YazerM,OlssonM,PalcicMOlssonPalcic(2006)."ThecisABbloodgroupphenotype:fundamentallessons
inglycobiology".TransfusMedRev20(3):20717.doi:10.1016/j.tmrv.2006.03.002
(https://dx.doi.org/10.1016%2Fj.tmrv.2006.03.002).PMID16787828
(https://www.ncbi.nlm.nih.gov/pubmed/16787828).
22. "ABOinheritancepatterns"(http://www.transfusion.com.au/?q=node/77).Inheritancepatternsofbloodgroups.
AustralianRedCrossBloodService.Retrieved30October2013.
23. "ABObloodgroupsystem"(http://abobloodtypes.webnode.com/).Abobloodtypes.webnode.com.Retrieved
20150202.
24. Citedin"Misattributedpaternityratesandnonpaternityrates"
(http://www.childsupportanalysis.co.uk/analysis_and_opinion/choices_and_behaviours/misattributed_paternity.ht
m).
25. "Understandinggenetics"(http://genetics.thetech.org/ask/ask221).Stanford.
26. BloodGroupASuptypes
(https://web.archive.org/web/20080802232301/http://www.owenfoundation.com/Health_Science/Blood_Group_A_
Subtypes.html),TheOwenFoundation.Retrieved1July2008.
27. SeltsamA,HallenslebenM,KollmannA,BlasczykR(2003)."Thenatureofdiversityanddiversificationatthe
ABOlocus".Blood102(8):303542.doi:10.1182/blood2003030955(https://dx.doi.org/10.1182%2Fblood
2003030955).PMID12829588(https://www.ncbi.nlm.nih.gov/pubmed/12829588).
28. OgasawaraKBannaiMSaitouNetal.(1996)."ExtensivepolymorphismofABObloodgroupgene:three
majorlineagesoftheallelesforthecommonABOphenotypes".HumanGenetics97(6):77783.
doi:10.1007/BF02346189(https://dx.doi.org/10.1007%2FBF02346189).PMID8641696
(https://www.ncbi.nlm.nih.gov/pubmed/8641696).
29. Shastry,SBhat,S(October2010)."ImbalanceinA2andA2BphenotypefrequencyofABOgroupinSouth
India"(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2957492).BloodTransfusion8(4):267270.
doi:10.2450/2010.014709(https://dx.doi.org/10.2450%2F2010.014709).PMC2957492
(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2957492).PMID20967168
(https://www.ncbi.nlm.nih.gov/pubmed/20967168).
30. Potts,WTW(1979)."HistoryandBloodGroupsintheBritishIsles".InSawyerPH.EnglishMedieval
Settlement.St.Martin'sPress.ISBN0713162570.
31. CsertiCM,DzikWH(2007)TheABObloodgroupsystemandPlasmodiumfalciparummalaria.Blood110(7)
22502258
32. Calafell,Francescetal.(September2008)."EvolutionarydynamicsofthehumanABOgene"
(http://www.springerlink.com/content/yv4072vu67mv1166/fulltext.html).HumanGenetics124(2):123135.
doi:10.1007/s0043900805308(https://dx.doi.org/10.1007%2Fs0043900805308).PMID18629539
(https://www.ncbi.nlm.nih.gov/pubmed/18629539).Retrieved24September2008.
33. LettertotheEditor:"Natural"VersusRegularAntibodies(http://resources.metapress.com/pdfpreview.axd?
code=l2206u8gj73x3251&size=largest)JournalTheProteinJournalPublisherSpringerNetherlandsISSN1572
3887(Print)15734943(Online)IssueVolume23,Number6/August,2004CategoryLettertotheEditorDOI
10.1023/B:JOPC.0000039625.56296.6ePage357SubjectGroupChemistryandMaterialsScienceOnlineDate
Friday,7January2005
34. Arendrup,MHansenJEClausenHNielsenCMathiesenLRNielsenJO(April1991)."Antibodytohisto
bloodgroupAantigenneutralizesHIVproducedbylymphocytesfrombloodgroupAdonorsbutnotfromblood
groupBorOdonors".AIDS5(4):4414.doi:10.1097/0000203019910400000014
(https://dx.doi.org/10.1097%2F0000203019910400000014).PMID1711864
(https://www.ncbi.nlm.nih.gov/pubmed/1711864).
35. Neil,SJMcKnightAGustafssonKWeissRA(20050615)."HIV1incorporatesABOhistobloodgroup
antigensthatsensitizevirionstocomplementmediatedinactivation"
(http://bloodjournal.hematologylibrary.org/cgi/content/full/105/12/4693).Blood105(12):46939.
doi:10.1182/blood2004114267(https://dx.doi.org/10.1182%2Fblood2004114267).PMID15728127
(https://www.ncbi.nlm.nih.gov/pubmed/15728127).
36. SeymourRM,AllanMJ,PomiankowskiA,andGustafssonK(2004)EvolutionoftheHumanABO
PolymorphismbyTwoComplementarySelectivePressures.Proceedings:BiologicalSciences271:10651072.
37. Reid,MEMohandas,N(2004)."Redbloodcellbloodgroupantigens:structureandfunction.".Seminarsin
hematology41(2):93117.doi:10.1053/j.seminhematol.2004.01.001
http://en.wikipedia.org/wiki/ABO_blood_group_system

11/14

5/12/2015

38.

39.

40.

41.

42.
43.

44.

45.

46.

47.

48.

ABObloodgroupsystemWikipedia,thefreeencyclopedia

(https://dx.doi.org/10.1053%2Fj.seminhematol.2004.01.001).PMID15071789
(https://www.ncbi.nlm.nih.gov/pubmed/15071789).
Mohandas,NNarla,A(2005)."Bloodgroupantigensinhealthanddisease.".CurrentOpinioninHematology
12(2):13540.doi:10.1097/01.moh.0000153000.09585.79
(https://dx.doi.org/10.1097%2F01.moh.0000153000.09585.79).PMID15725904
(https://www.ncbi.nlm.nih.gov/pubmed/15725904).
Sarode,RGoldsteinJSussmanIINagelRLTsaiHM(June2000)."RoleofAandBbloodgroupantigensin
theexpressionofadhesiveactivityofvonWillebrandfactor".BrJHaematol.109(4):85764.
doi:10.1046/j.13652141.2000.02113.x(https://dx.doi.org/10.1046%2Fj.13652141.2000.02113.x).
PMID10929042(https://www.ncbi.nlm.nih.gov/pubmed/10929042).
O'Donnell,JLaffanMA(August2001)."TherelationshipbetweenABOhistobloodgroup,factorVIIIandvon
Willebrandfactor".TransfusMed.11(4):34351.doi:10.1046/j.13653148.2001.00315.x
(https://dx.doi.org/10.1046%2Fj.13653148.2001.00315.x).PMID11532189
(https://www.ncbi.nlm.nih.gov/pubmed/11532189).
O'Donnell,JBoultonFEManningRALaffanMA(20020201)."AmountofHantigenexpressedon
circulatingvonWillebrandfactorismodifiedbyABObloodgroupgenotypeandisamajordeterminantofplasma
vonWillebrandfactorantigenlevels"(http://atvb.ahajournals.org/cgi/content/full/22/2/335).ArteriosclerThromb
VascBiol.(AmericanHeartAssociation,Inc.)22(2):33541.doi:10.1161/hq0202.103997
(https://dx.doi.org/10.1161%2Fhq0202.103997).PMID11834538
(https://www.ncbi.nlm.nih.gov/pubmed/11834538).
Gill,JCEndresBrooksJBauerPJMarksWJMontgomeryRR(June1987)."TheeffectofABObloodgroup
onthediagnosisofvonWillebranddisease"(http://www.bloodjournal.org/cgi/content/abstract/69/6/1691)
(ABSTRACT).Blood69(6):16915.PMID3495304(https://www.ncbi.nlm.nih.gov/pubmed/3495304).
Shima,MFujimuraYetal.(1995)."ABObloodgroupgenotypeandplasmavonWillebrandfactorinnormal
individuals".VoxSang68(4):23640.doi:10.1111/j.14230410.1995.tb02579.x
(https://dx.doi.org/10.1111%2Fj.14230410.1995.tb02579.x).PMID7660643
(https://www.ncbi.nlm.nih.gov/pubmed/7660643).
Bowen,DJCollinsPWetal.(March2005)."Theprevalenceofthecysteine1584variantofvonWillebrand
factorisincreasedintype1vonWillebranddisease:cosegregationwithincreasedsusceptibilitytoADAMTS13
proteolysisbutnotclinicalphenotype".BrJHaematol(BlackwellSynergy)128(6):8306.doi:10.1111/j.1365
2141.2005.05375.x(https://dx.doi.org/10.1111%2Fj.13652141.2005.05375.x).PMID15755288
(https://www.ncbi.nlm.nih.gov/pubmed/15755288).
BongersTdeMaatMvanGoorMetal.(2006)."HighvonWillebrandfactorlevelsincreasetheriskoffirst
ischemicstroke:influenceofADAMTS13,inflammation,andgeneticvariability".Stroke37(11):26727.
doi:10.1161/01.STR.0000244767.39962.f7(https://dx.doi.org/10.1161%2F01.STR.0000244767.39962.f7).
PMID16990571(https://www.ncbi.nlm.nih.gov/pubmed/16990571).
XieJ,QureshiAA,LiY,HanJQureshiLiHan(2010)."ABOBloodGroupandIncidenceofSkinCancer"
(http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0011972).PLoSONE5(8):e11972.
Bibcode:2010PLoSO...511972X(http://adsabs.harvard.edu/abs/2010PLoSO...511972X).
doi:10.1371/journal.pone.0011972(https://dx.doi.org/10.1371%2Fjournal.pone.0011972).
Wolpin,B.M.Kraft,P.Gross,M.Helzlsouer,K.BuenoDeMesquita,H.B.Steplowski,E.Stolzenberg
Solomon,R.Z.Arslan,A.A.Jacobs,E.J.Lacroix,A.Petersen,G.Zheng,W.Albanes,D.Allen,N.E.
Amundadottir,L.Anderson,G.BoutronRuault,M.C.Buring,J.E.Canzian,F.Chanock,S.J.Clipp,S.
Gaziano,J.M.Giovannucci,E.L.Hallmans,G.Hankinson,S.E.Hoover,R.N.Hunter,D.J.Hutchinson,
A.Jacobs,K.Kooperberg,C.(2010)."PancreaticCancerRiskandABOBloodGroupAlleles:Resultsfromthe
PancreaticCancerCohortConsortium"(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2943735).Cancer
Research70(3):10151023.doi:10.1158/00085472.CAN092993(https://dx.doi.org/10.1158%2F0008
5472.CAN092993).PMC2943735(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2943735).
PMID20103627(https://www.ncbi.nlm.nih.gov/pubmed/20103627).
Amundadottir,L.Kraft,P.StolzenbergSolomon,R.Z.Fuchs,C.S.Petersen,G.M.Arslan,A.A.Bueno
DeMesquita,H.B.Gross,M.Helzlsouer,K.Jacobs,E.J.Lacroix,A.Zheng,W.Albanes,D.Bamlet,W.
Berg,C.D.Berrino,F.Bingham,S.Buring,J.E.Bracci,P.M.Canzian,F.ClavelChapelon,F.O.Clipp,
S.Cotterchio,M.DeAndrade,M.Duell,E.J.FoxJr,J.W.Gallinger,S.Gaziano,J.M.Giovannucci,E.
L.Goggins,M.(2009)."GenomewideassociationstudyidentifiesvariantsintheABOlocusassociatedwith

http://en.wikipedia.org/wiki/ABO_blood_group_system

12/14

5/12/2015

49.

50.

51.

52.

53.
54.
55.

ABObloodgroupsystemWikipedia,thefreeencyclopedia

susceptibilitytopancreaticcancer"(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2839871).NatureGenetics
41(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).
Gates,MAWolpin,BMCramer,DWHankinson,SETworoger,SS(2010)."ABObloodgroupandincidence
ofepithelialovariancancer"(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2946962).InternationalJournalof
Cancer.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).
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).
Glass,RogerHolmgrenJ,HaleyC,KhanM.R.,SVENERHOLMM,STOLLB,HOSSAINK,BLACKB,
YunusM,BaruaD(June1985)."PredispositionforcholeraofindividualswithObloodgroup.Possible
evolutionarysignificance"(http://eutils.ncbi.nlm.nih.gov/entrez/eutils/elink.fcgi?
dbfrom=pubmed&id=4014172&retmode=ref&cmd=prlinks).AmericanJournalofEpidemiology121(6):791
796.PMID4014172(https://www.ncbi.nlm.nih.gov/pubmed/4014172).Retrieved20August2012.
AmericanRedCross,SouthernCaliforniaBloodServicesRegion(n.d.)."AnswerstoCommonlyAsked
QuestionsAboutBloodandBloodBanking"(http://www.socalredcross.org/pdf/BloodThe.pdf)(PDF).Blood:the
Basics:4.Archived
(http://web.archive.org/web/20071129152741/http://www.socalredcross.org/pdf/BloodThe.pdf)(PDF)fromthe
originalon29November2007.Retrieved16November2007.
SungIlRyu,YoungWooSohn(2007),AReviewofSociocultural,Behavioral,BiochemicalAnalysesonABO
BloodGroupsTypology,TheKoreanJournalofSocialandPersonalityPsychology.
DonnaK.Hobgood(2011),Personalitytraitsofaggressionsubmissivenessandperfectionismassociatewith
ABObloodgroupsthroughcatecholamineactivities
(http://www.sciencedirect.com/science/article/pii/S0306987711002106),MedicalHypotheses,77(2):294300.
Klein,HarveyG(7March2005)."WhyDoPeopleHaveDifferentBloodTypes?"
(http://www.sciam.com/article.cfm?id=whydopeoplehavediffer).ScientificAmerican.Retrieved16November
2007.

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)at
BGMUTBloodGroupAntigenGeneMutationDatabaseatNCBI,NIH
ABObloodgroups,antibodiesandantigensexplained(https://www.youtube.com/watch?
v=u7DxZmLWDII)YouTubeeducationalvideo
http://en.wikipedia.org/wiki/ABO_blood_group_system

13/14

5/12/2015

ABObloodgroupsystemWikipedia,thefreeencyclopedia

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"
Categories: Bloodantigensystems
Thispagewaslastmodifiedon9May2015,at17:20.
TextisavailableundertheCreativeCommonsAttributionShareAlikeLicenseadditionaltermsmay
apply.Byusingthissite,youagreetotheTermsofUseandPrivacyPolicy.Wikipediaisa
registeredtrademarkoftheWikimediaFoundation,Inc.,anonprofitorganization.

http://en.wikipedia.org/wiki/ABO_blood_group_system

14/14