ABO Blood Group System - Wikipedia, The Free Encyclopedia

5/12/2015 ABObloodgroupsystemWikipedia,thefreeencyclopedia

http://en.wikipedia.org/wiki/ABO_blood_group_system 1/14

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



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



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



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]



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]



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



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



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



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]



Seealso

ArtificialbloodBloodtransfusionBombayphenotypeCisABKiddbloodgroupRhbloodgroupsystem

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33. LettertotheEditor:"Natural"VersusRegularAntibodies(http://resources.metapress.com/pdfpreview.axd?code=l2206u8gj73x3251&size=largest)JournalTheProteinJournalPublisherSpringerNetherlandsISSN15723887(Print)15734943(Online)IssueVolume23,Number6/August,2004CategoryLettertotheEditorDOI10.1023/B:JOPC.0000039625.56296.6ePage357SubjectGroupChemistryandMaterialsScienceOnlineDateFriday,7January2005

34. Arendrup,MHansenJEClausenHNielsenCMathiesenLRNielsenJO(April1991)."AntibodytohistobloodgroupAantigenneutralizesHIVproducedbylymphocytesfrombloodgroupAdonorsbutnotfrombloodgroupBorOdonors".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)."HIV1incorporatesABOhistobloodgroupantigensthatsensitizevirionstocomplementmediatedinactivation"(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)EvolutionoftheHumanABOPolymorphismbyTwoComplementarySelectivePressures.Proceedings:BiologicalSciences271:10651072.

37. Reid,MEMohandas,N(2004)."Redbloodcellbloodgroupantigens:structureandfunction.".Seminarsinhematology41(2):93117.doi:10.1053/j.seminhematol.2004.01.001



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38. Mohandas,NNarla,A(2005)."Bloodgroupantigensinhealthanddisease.".CurrentOpinioninHematology12(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).

39. Sarode,RGoldsteinJSussmanIINagelRLTsaiHM(June2000)."RoleofAandBbloodgroupantigensintheexpressionofadhesiveactivityofvonWillebrandfactor".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).

40. O'Donnell,JLaffanMA(August2001)."TherelationshipbetweenABOhistobloodgroup,factorVIIIandvonWillebrandfactor".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).

41. O'Donnell,JBoultonFEManningRALaffanMA(20020201)."AmountofHantigenexpressedoncirculatingvonWillebrandfactorismodifiedbyABObloodgroupgenotypeandisamajordeterminantofplasmavonWillebrandfactorantigenlevels"(http://atvb.ahajournals.org/cgi/content/full/22/2/335).ArteriosclerThrombVascBiol.(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).

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44. Bowen,DJCollinsPWetal.(March2005)."Theprevalenceofthecysteine1584variantofvonWillebrandfactorisincreasedintype1vonWillebranddisease:cosegregationwithincreasedsusceptibilitytoADAMTS13proteolysisbutnotclinicalphenotype".BrJHaematol(BlackwellSynergy)128(6):8306.doi:10.1111/j.13652141.2005.05375.x(https://dx.doi.org/10.1111%2Fj.13652141.2005.05375.x).PMID15755288(https://www.ncbi.nlm.nih.gov/pubmed/15755288).

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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).

51. Glass,RogerHolmgrenJ,HaleyC,KhanM.R.,SVENERHOLMM,STOLLB,HOSSAINK,BLACKB,YunusM,BaruaD(June1985)."PredispositionforcholeraofindividualswithObloodgroup.Possibleevolutionarysignificance"(http://eutils.ncbi.nlm.nih.gov/entrez/eutils/elink.fcgi?dbfrom=pubmed&id=4014172&retmode=ref&cmd=prlinks).AmericanJournalofEpidemiology121(6):791796.PMID4014172(https://www.ncbi.nlm.nih.gov/pubmed/4014172).Retrieved20August2012.

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.



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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