abo blood group system - wikipedia, the free encyclopedia

ABO blood group antigens present on red blood cells and IgM

antibodies present in the serum

ABO blood group systemFrom Wikipedia, the free encyclopedia

The ABO blood group system is the mostimportant blood type system (or blood groupsystem) in human blood transfusion. Theassociated anti-A antibodies and anti-Bantibodies are usually IgM antibodies, whichare usually produced in the first years of lifeby sensitization to environmental substancessuch as food, bacteria, and viruses. ABOblood types are also present in someanimals, for example apes such as

chimpanzees, bonobos, and gorillas.[1]

Contents

1 History of discoveries2 ABO antigens3 Serology

3.1 Origin theories

4 Nonantigen biology5 Transfusion reactions6 ABO hemolytic disease of thenewborn7 Inheritance8 Distribution and evolutionaryhistory

8.1 ABO and Rhdistribution bycountry

9 Association with vonWillebrand factor10 Disease association11 Subgroups

11.1 A1 and A2

12 Bombay phenotype13 Nomenclature in Europeand former USSR14 Examples of ABO andRhesus D slide testing method15 Universal blood createdfrom other types, and artificialblood16 Conjectures17 See also

ABO blood group system - Wikipedia, the free encyclopedia http://en.wikipedia.org/wiki/ABO_blood_group_system

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Diagram showing the carbohydrate chains that

determine the ABO blood group

18 References19 Further reading20 External links

History of discoveries

The ABO blood group system is widely credited to have been discovered by the Austrian scientist Karl

Landsteiner, who found three different blood types in 1900;[2] he was awarded the Nobel Prize in Physiology orMedicine in 1930 for his work. Due to inadequate communication at the time it was subsequently found that

Czech serologist Jan Janský had independently pioneered the classification of human blood into four groups,[3]

but Landsteiner's independent discovery had been accepted by the scientific world while Janský remained inrelative obscurity. Janský's classification is however still used in Russia and states of former USSR (see below).

In America, Moss published his own (very similar) work in 1910.[4]

Landsteiner described A, B, and O; Alfred von Decastello and Adriano Sturli discovered the fourth type, AB, in

1902.[5] Ludwik Hirszfeld and E. von Dungern discovered the heritability of ABO blood groups in 1910–11,with Felix Bernstein demonstrating the correct blood group inheritance pattern of multiple alleles at one locus in

1924.[6] Watkins and Morgan, in England, discovered that the ABO epitopes were conferred by sugars, to be

specific, N-acetylgalactosamine for the A-type and galactose for the B-type.[7][8][9] After much publishedliterature claiming that the ABH substances were all attached to glycosphingolipids, Laine's group (1988) found

that the band 3 protein expressed a long polylactosamine chain[10] that contains the major portion of the ABH

substances attached.[11] Later, Yamamoto's group showed the precise glycosyl transferase set that confers the

A, B and O epitopes.[12]

ABO antigens

The H antigen is an essential precursor to the ABO bloodgroup antigens. The H locus is located on chromosome 19.It contains 3 exons that span more than 5 kb of genomicDNA, and it encodes a fucosyltransferase that producesthe H antigen on RBCs. The H antigen is a carbohydratesequence with carbohydrates linked mainly to protein(with a minor fraction attached to ceramide moiety). Itconsists of a chain of β-D-galactose, β-D-N-acetylglucosamine, β-D-galactose, and 2-linked,α-L-fucose, the chain being attached to the protein orceramide.

The ABO locus is located on chromosome 9. It contains 7exons that span more than 18 kb of genomic DNA. Exon 7is the largest and contains most of the coding sequence.The ABO locus has three main alleleic forms: A, B, and O.The A allele encodes a glycosyltransferase that bondsα-N-acetylgalactosamine to D-galactose end of H antigen, producing the A antigen. The B allele encodes aglycosyltransferase that joins α-D-galactose bonded to D-galactose end of H antigen, creating the B antigen.


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In case of O allele, the exon 6 contains a deletion that results in a loss of enzymatic activity. The O allele differsfrom the A allele by deletion of only one nucleotide – guanine at position 261. The deletion causes a frameshift,and results in premature termination of translation, and thus, degradation of the mRNA. This results in H antigenremaining unchanged in case of O groups.

The majority of the ABO antigens are expressed on the ends of long polylactosamine chains attached mainly toband 3 protein, the anion exchange protein of the RBC membrane, and a minority of the epitopes are expressedon neutral glycosphingolipids.

Serology

Anti-A and anti-B antibodies (called isohaemagglutinins), which are not present in the newborn, appear in thefirst years of life. They are isoantibodies, that is, they are produced by an individual against antigens producedby members of the same species (isoantigens). Anti-A and anti-B antibodies are usually IgM type, which are notable to pass through the placenta to the fetal blood circulation. O-type individuals can produce IgG-type ABOantibodies.

Origin theories

It is possible that food and environmental antigens (bacterial, viral, or plant antigens) have epitopes similarenough to A and B glycoprotein antigens. The antibodies created against these environmental antigens in thefirst years of life can cross-react with ABO-incompatible red blood cells that it comes in contact with duringblood transfusion later in life. Anti-A antibodies are hypothesized to originate from immune response towardsinfluenza virus, whose epitopes are similar enough to the α-D-N-galactosamine on the A glycoprotein to be ableto elicit a cross-reaction. Anti-B antibodies are hypothesized to originate from antibodies produced against

Gram-negative bacteria, such as E. coli, cross-reacting with the α-D-galactose on the B glycoprotein.[13]

The "Light in the Dark theory" (DelNagro, 1998) suggests that, when budding viruses acquire host cellmembranes from one human patient (in particular, from the lung and mucosal epithelium where they are highlyexpressed), they also take along ABO blood antigens from those membranes, and may carry them intosecondary recipients where these antigens can elicit a host immune response against these non-self foreignblood antigens. These viral-carried human blood antigens may be responsible for priming newborns intoproducing neutralizing antibodies against foreign blood antigens. Support for this theory has come to light inrecent experiments with HIV. HIV can be neutralized in in vitro experiments using antibodies against blood

group antigens specifically expressed on the HIV-producing cell lines.[14][15]

The "Light in the Dark theory" suggests a novel evolutionary hypothesis: there is true communal immunity,which has developed to reduce the inter-transmissibility of viruses within a population. It suggests thatindividuals in a population supply and make a diversity of unique antigenic moieties so as to keep the populationas a whole more resistant to infection. A system set up ideally to work with variable recessive alleles.[citation needed]

However, it is more likely that the force driving evolution of allele diversity is simply negative frequency-dependent selection; cells with rare variants of membrane antigens are more easily distinguished by the immunesystem from pathogens carrying antigens from other hosts. Thus, individuals possessing rare types are betterequipped to detect pathogens. The high within-population diversity observed in human populations would, then,

be a consequence of natural selection on individuals[16]

Nonantigen biology


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The carbohydrate molecules on the surfaces of red blood cells have roles in cell membrane integrity, celladhesion, membrane transportation of molecules, and acting as receptors for extracellular ligands, and enzymes.

ABO antigens are found having similar roles on epithelial cells as well as red blood cells.[17][18]

Transfusion reactions

Due to the presence of isoantibodies against non-self blood group antigens, individuals of type A blood groupimmediately raise anti-B antibodies against B-blood group RBCs if transfused with blood from B group. Theanti-B antibodies bind to B antigens on RBCs and cause complement-mediated lysis of the RBCs. The samehappens for B and O groups (which raises both anti-A and anti-B antibodies). However, only blood group ABdoes not have anti-A and anti-B isoantibodies. This is because both A and B-antigens are present on the RBCsand are both self-antigens, hence they can receive blood from all groups and are universal recipient.

As far as transfusion compatibility is concerned, it is not strictly as simple as matching A, B, and O groups. Inother words, no individual will ever receive a blood transfusion based on the ABO system alone. The rhesusfactor must also be considered. Together, the rhesus factor and ABO grouping are the two most importantcompatibility factors to consider. An individual may be Rh+ or Rh-. In simpler terms, if an individual is bloodtype A and positive for the rhesus factor, then he or she is deemed "A+".

ABO and Rh blood type donation showing matches between donor and recipient types

Recipients Donors

O+ A+ B+ AB+ O- ** A- B- AB-

O+ • •

A+ • • • •

B+ • • • •

AB+ * • • • • • • • •

O- •

A- • •

B- • •

AB- • • • •

* Type AB+ is the universal recipient: Although those with AB blood type may be referred to as universalrecipients, in actuality, type AB+ blood is that of the universal recipient, whereas type AB- is not. This is animportant distinction to make.

** Because A-, A+, B-, B+, AB-, AB+, O- and O+ individuals can all receive blood from donors of type O-blood, an individual with type O- blood is deemed universal donor. In similar manner, O+ is not theuniversal donor blood type.

One caveat to this axiom of 'universal donor' is that this applies to packed RBCs, and not to whole bloodproducts. Using the first table, type O carries anti-A and anti-B antibodies in the serum. To transfuse a type A,B, or AB recipient with type O whole blood would produce a hemolytic transfusion reaction due to theantibodies found in the serum of whole blood.

No antibodies are formed against the H antigen, except in those individuals with the Bombay phenotype.


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A and B are codominant, giving the AB

phenotype.

In ABH secretors, ABH antigens are secreted by most mucus-producing cells of the body interfacing with the

environment, including lung, skin, liver, pancreas, stomach, intestines, ovaries, and prostate.[19]

ABO hemolytic disease of the newborn

Main article: Hemolytic disease of the newborn (ABO)

ABO blood group incompatibilities between the mother and child does not usually cause hemolytic disease ofthe newborn (HDN) because antibodies to the ABO blood groups are usually of the IgM type, which do notcross the placenta; however, in an O-type mother, IgG ABO antibodies are produced and the baby can developABO hemolytic disease of the newborn.

Inheritance

Blood groups are inherited from both parents. The ABO bloodtype is controlled by a single gene (the ABO gene) with three

alleles: i, IA, and IB. The gene encodes aglycosyltransferase—that is, an enzyme that modifies thecarbohydrate content of the red blood cell antigens. The gene islocated on the long arm of the ninth chromosome (9q34).

The IA allele gives type A, IB gives type B, and i gives type O.

As both IA and IB are dominant over i, only ii people have type

O blood. Individuals with IAIA or IAi have type A blood, and

individuals with IBIB or IBi have type B. IAIB people have bothphenotypes, because A and B express a special dominancerelationship: codominance, which means that type A and Bparents can have an AB child. A type A and a type B couplecan also have a type O child if they are both heterozygous

(IBi,IAi) The cis-AB phenotype has a single enzyme that createsboth A and B antigens. The resulting red blood cells do notusually express A or B antigen at the same level that would be

expected on common group A1 or B red blood cells, which can help solve the problem of an apparently

genetically impossible blood group.[20]

Distribution and evolutionary history

The distribution of the blood groups A, B, O and AB varies across the world according to the population. Thereare also variations in blood type distribution within human subpopulations.

In the UK, the distribution of blood type frequencies through the population still shows some correlation to thedistribution of placenames and to the successive invasions and migrations including Vikings, Danes, Saxons,

Celts, and Normans who contributed the morphemes to the placenames and the genes to the population.[21]

There are six common alleles in white individuals of the ABO gene that produce one's blood type:[22][23]


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Frequency of O group in indigenous populations around the

world

A

A101 (A1)A201 (A2)

B

B101 (B1)

O

O01 (O1)O02 (O1v)O03 (O2)

Many rare variants of these alleles have been found in human populations around the world.

Some evolutionary biologists theorize that the IA allele evolved earliest, followed by O (by the deletion of a

single nucleotide, shifting the reading frame) and then IB.[citation needed] This chronology accounts for thepercentage of people worldwide with each blood type. It is consistent with the accepted patterns of earlypopulation movements and varying prevalent blood types in different parts of the world: for instance, B is verycommon in populations of Asian descent, but rare in ones of Western European descent. Another theory statesthat there are four main lineages of the ABO gene and that mutations creating type O have occurred at least

three times in humans.[24] From oldest to youngest, these lineages comprise the following alleles:A101/A201/O09, B101, O02 and O01. The continued presence of the O alleles is hypothesized to be the result

of balancing selection.[24] Both theories contradict the previously held theory that type O blood evolvedearliest.

ABO and Rh distribution by country

ABO and Rh blood type distribution by country (population averages)

Country Population[25] O+ A+ B+ AB+ O- A- B- AB-

Australia[26] 21,262,641 40.0% 31.0% 8.0% 2.0% 9.0% 7.0% 2.0% 1.0%

Austria[27] 8,210,281 30.0% 33.0% 12.0% 6.0% 7.0% 8.0% 3.0% 1.0%

Belgium[28] 10,414,336 38.0% 34.0% 8.5% 4.1% 7.0% 6.0% 1.5% 0.8%

Brazil[29] 198,739,269 36.0% 34.0% 8.0% 2.5% 9.0% 8.0% 2.0% 0.5%

Canada[30] 33,487,208 39.0% 36.0% 7.6% 2.5% 7.0% 6.0% 1.4% 0.5%

Czech Republic[31] 10,532,770 27.0% 36.0% 15.0% 7.0% 5.0% 6.0% 3.0% 1.0%


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Denmark[32] 5,500,510 35.0% 37.0% 8.0% 4.0% 6.0% 7.0% 2.0% 1.0%

Estonia[33] 1,299,371 30.0% 31.0% 20.0% 6.0% 4.5% 4.5% 3.0% 1.0%

Finland[34] 5,250,275 27.0% 38.0% 15.0% 7.0% 4.0% 6.0% 2.0% 1.0%

France[35] 62,150,775 36.0% 37.0% 9.0% 3.0% 6.0% 7.0% 1.0% 1.0%

Germany[36] 82,329,758 35.0% 37.0% 9.0% 4.0% 6.0% 6.0% 2.0% 1.0%

Hong Kong SAR[37] 7,055,071 40.0% 26.0% 27.0% 7.0% 0.3% 0.2% 0.1% 0.1%

Iceland[38] 306,694 47.6% 26.4% 9.3% 1.6% 8.4% 4.6% 1.7% 0.4%

India[39] 1,166,079,217 36.5% 22.1% 30.9% 6.4% 2.0% 0.8% 1.1% 0.2%

Ireland[40] 4,203,200 47.0% 26.0% 9.0% 2.0% 8.0% 5.0% 2.0% 1.0%

Israel[41] 7,233,701 32.0% 34.0% 17.0% 7.0% 3.0% 4.0% 2.0% 1.0%

Netherlands[42] 16,715,999 39.5% 35.0% 6.7% 2.5% 7.5% 7.0% 1.3% 0.5%

New Zealand[43] 4,213,418 38.0% 32.0% 9.0% 3.0% 9.0% 6.0% 2.0% 1.0%

Norway[44] 4,660,539 34.0% 42.5% 6.8% 3.4% 6.0% 7.5% 1.2% 0.6%

Poland[45] 38,482,919 31.0% 32.0% 15.0% 7.0% 6.0% 6.0% 2.0% 1.0%

Portugal[46] 10,707,924 36.2% 39.8% 6.6% 2.9% 6.0% 6.6% 1.1% 0.5%

Saudi Arabia[47] 28,686,633 48.0% 24.0% 17.0% 4.0% 4.0% 2.0% 1.0% 0.3%

South Africa[48] 49,320,000 39.0% 32.0% 12.0% 3.0% 7.0% 5.0% 2.0% 1.0%

Spain[49] 40,525,002 36.0% 34.0% 8.0% 2.5% 9.0% 8.0% 2.0% 0.5%

Sweden[50] 9,059,651 32.0% 37.0% 10.0% 5.0% 6.0% 7.0% 2.0% 1.0%

Turkey[51] 76,805,524 29.8% 37.8% 14.2% 7.2% 3.9% 4.7% 1.6% 0.8%

United Kingdom[52] 61,113,205 37.0% 35.0% 8.0% 3.0% 7.0% 7.0% 2.0% 1.0%

United States[53] 307,212,123 37.4% 35.7% 8.5% 3.4% 6.6% 6.3% 1.5% 0.6%

Weighted mean 2,261,025,244 36.4% 28.3% 20.6% 5.1% 4.3% 3.5% 1.4% 0.5%

Racial & Ethnic Distribution of ABO (without Rh) Blood Types[54]

(This table has more entries than the table above but does not distinguish between Rh types.)

PEOPLE GROUP O (%) A (%) B (%)AB (%)

Aborigines 61 39 0 0

Abyssinians 43 27 25 5

Ainu (Japan) 17 32 32 18


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Albanians 38 43 13 6

Grand Andamanese 9 60 23 9

Arabs 34 31 29 6

Armenians 31 50 13 6

Asian (in USA - General) 40 28 27 5

Austrians 36 44 13 6

Bantus 46 30 19 5

Basques 51 44 4 1

Belgians 47 42 8 3

Blackfoot (N. Am. Indian) 17 82 0 1

Bororo (Brazil) 100 0 0 0

Brazilians 47 41 9 3

Bulgarians 32 44 15 8

Burmese 36 24 33 7

Buryats (Siberia) 33 21 38 8

Bushmen 56 34 9 2

Chinese-Canton 46 23 25 6

Chinese-Peking 29 27 32 13

Chuvash 30 29 33 7

Czechs 30 44 18 9

Danes 41 44 11 4

Dutch 45 43 9 3

Egyptians 33 36 24 8

English 47 42 9 3

Eskimos (Alaska) 38 44 13 5

Eskimos (Greenland) 54 36 23 8

Estonians 34 36 23 8

Fijians 44 34 17 6

Finns 34 41 18 7

French 43 47 7 3

Georgians 46 37 12 4

Germans 41 43 11 5

Greeks 40 42 14 5

Gypsies (Hungary) 29 27 35 10


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Hawaiians 37 61 2 1

Hindus (Bombay) 32 29 28 11

Hungarians 36 43 16 5

Icelanders 56 32 10 3

Indians (India - General) 37 22 33 7

Indians (USA - General) 79 16 4 1

Irish 52 35 10 3

Italians (Milan) 46 41 11 3

Japanese 30 38 22 10

Jews (Germany) 42 41 12 5

Jews (Poland) 33 41 18 8

Kalmuks 26 23 41 11

Kikuyu (Kenya) 60 19 20 1

Koreans 28 32 31 10

Lapps 29 63 4 4

Latvians 32 37 24 7

Lithuanians 40 34 20 6

Malaysians 62 18 20 0

Maori 46 54 1 0

Mayas 98 1 1 1

Moros 64 16 20 0

Navajo (N. Am. Indian) 73 27 0 0

Nicobarese (Nicobars) 74 9 15 1

Norwegians 39 50 8 4

Papuas (New Guinea) 41 27 23 9

Persians 38 33 22 7

Peru (Indians) 100 0 0 0

Filipinos 45 22 27 6

Poles 33 39 20 9

Portuguese 35 53 8 4

Romanians 34 41 19 6

Russians 33 36 23 8

Sardinians 50 26 19 5

Scots 51 34 12 3


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Serbians 38 42 16 5

Shompen (Nicobars) 100 0 0 0

Slovaks 42 37 16 5

South Africans 45 40 11 4

Spanish 38 47 10 5

Sudanese 62 16 21 0

Swedes 38 47 10 5

Swiss 40 50 7 3

Tartars 28 30 29 13

Thais 37 22 33 8

Turks 43 34 18 6

Ukrainians 37 40 18 6

USA (US blacks) 49 27 20 4

USA (US whites) 45 40 11 4

Vietnamese 42 22 30 5

Mean 43.91 34.80 16.55 5.14

Standard deviation 16.87 13.80 9.97 3.41

Blood group B has its highest frequency in Northern India and neighboring Central Asia, and its incidence

diminishes both towards the west and the east, falling to single digit percentages in Spain.[55][56] It is believed tohave been entirely absent from Native American and Australian Aboriginal populations prior to the arrival of

Europeans in those areas.[56][57]

Blood group A is associated with high frequencies in Europe, especially in Scandinavia and Central Europe,although its highest frequencies occur in some Australian Aborigine populations and the Blackfoot Indians of

Montana.[58][59]

Association with von Willebrand factor

The ABO antigen is also expressed on the von Willebrand factor (vWF) glycoprotein,[60] which participates in

hemostasis (control of bleeding). In fact, having type O blood predisposes to bleeding,[61] as 30% of the total

genetic variation observed in plasma vWF is explained by the effect of the ABO blood group,[62] andindividuals with group O blood normally have significantly lower plasma levels of vWF (and Factor VIII) than

do non-O individuals.[63][64] In addition, vWF is degraded more rapidly due to the higher prevalence of blood

group O with the Cys1584 variant of vWF (an amino acid polymorphism in VWF):[65] the gene for ADAMTS13(vWF-cleaving protease) maps to the ninth chromosome (9q34), the same locus as ABO blood type. Higherlevels of vWF are more common amongst people who have had ischaemic stroke (from blood clotting) for the

first time.[66] The results of this study found that the occurrence was not affected by ADAMTS13


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Ukraine uniform imprint B+

polymorphism, and the only significant genetic factor was the person's blood group.

Disease association

Compared to non-O group (A, AB, and B) individuals, O group individuals have a 14% reduced risk of

squamous cell carcinoma and 4% reduced risk of basal cell carcinoma.[67] It is also associated with a reduced

risk of pancreatic cancer.[68][69] The B antigen links with increased risk of ovarian cancer.[70] Gastric cancer

has reported to be more common in blood group A and least in group O.[71]

Subgroups

A1 and A2

The A blood type contains about twenty subgroups, of which A1 and A2 are the most common (over 99%). A1

makes up about 80% of all A-type blood, with A2 making up the rest.[72] These two subgroups areinterchangeable as far as transfusion is concerned, but complications can sometimes arise in rare cases when

typing the blood.[72]

Bombay phenotype

Main article: Hh antigen system

Individuals with the rare Bombay phenotype (hh) do not express antigen H on their red blood cells. As Hantigen serves as precursor for producing A and B antigens, the absence of H antigen means the individuals donot have A or B antigens as well (similar to O blood group). However, unlike O group, the H antigen is absent,hence the individuals produce isoantibodies to antigen H as well as to both A and B antigens. In case theyreceive blood from O blood group, the anti-H antibodies will bind to H antigen on RBC of donor blood anddestroy the RBCs by complement-mediated lysis. Therefore Bombay phenotype can receive blood only fromother hh donors (although they can donate as though they were type O).

Nomenclature in Europe and former USSR

In parts of Europe, the "O" in ABO blood type is substituted with "0"(zero), signifying the lack of A or B antigen. In the former USSR bloodtypes are referenced using numbers and Roman numerals instead ofletters. This is Janský's original classification of blood types. Itdesignates the blood types of humans as I, II, III, and IV, which are

elsewhere designated, respectively, as O, A, B, and AB.[73] Thedesignation A and B with reference to blood groups was proposed by Ludwik Hirszfeld.

Examples of ABO and Rhesus D slide testing method


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Blood group Opositive: neither anti-Anor anti-B haveagglutinated, butanti-Rh has

Result: Blood group Apositive: anti-A andanti-Rh haveagglutinated but anti-Bhas not.

In the slide testing method shown above, three drops of blood are placed on a glass slide with liquid reagents.Agglutination indicates the presence of blood group antigens in the blood.

Universal blood created from other types, and artificial blood

In April 2007, an international team of researchers announced in the journal Nature Biotechnology an

inexpensive and efficient way to convert types A, B, and AB blood into type O.[74] This is done by usingglycosidase enzymes from specific bacteria to strip the blood group antigens from red blood cells. The removalof A and B antigens still does not address the problem of the Rhesus blood group antigen on the blood cells ofRhesus positive individuals, and so blood from Rhesus negative donors must be used. Patient trials will beconducted before the method can be relied on in live situations.

Another approach to the blood antigen problem is the manufacture of artificial blood, which could act as asubstitute in emergencies. BBC (http://news.bbc.co.uk/2/hi/uk_news/england/north_yorkshire/6645923.stm) .

Conjectures

There are numerous popular conjectures surrounding ABO blood groups. These beliefs have existed since theABO blood groups were identified and can be found in different cultures throughout the world. For example,during the 1930s, connecting blood groups to personality types became popular in Japan and other areas of the

world.[75]

The popularity of Peter J. D'Adamo's book, Eat Right For Your Blood Type suggests that these conjectures

persist. This book claims that ABO blood type determines one's optimal diet.[76]

Additional myths include the idea that group A causes severe hangovers, group O is associated with perfectteeth, and those with blood group A2 have the highest IQs. Scientific evidence in support of these concepts is

nonexistent.[77]

See also

Cis AB


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^ Neil, SJ; McKnight A, Gustafsson K, Weiss RA(2005-06-15). "HIV-1 incorporates ABO histo-bloodgroup antigens that sensitize virions to complement-mediated inactivation"(http://bloodjournal.hematologylibrary.org/cgi/content/full/105/12/4693) . Blood 105 (12):4693–9. doi:10.1182/blood-2004-11-4267(http://dx.doi.org/10.1182%2Fblood-2004-11-4267) .PMID 15728127 (http://www.ncbi.nlm.nih.gov/pubmed/15728127) .http://bloodjournal.hematologylibrary.org/cgi/content/full/105/12/4693.

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^ Seymour RM, Allan MJ, Pomiankowski A, andGustafsson K (2004) Evolution of the Human ABOPolymorphism by Two Complementary SelectivePressures. Proceedings: Biological Sciences271:1065-1072.

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^ Reid, ME; Mohandas, N (2004). "Red blood cellblood group antigens: structure and function.".Seminars in hematology 41 (2): 93–117.doi:10.1053/j.seminhematol.2004.01.001(http://dx.doi.org/10.1053%2Fj.seminhematol.2004.01.001) .PMID 15071789 (http://www.ncbi.nlm.nih.gov/pubmed/15071789) .

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^ Mohandas, N; Narla, A (2005). "Blood groupantigens in health and disease.". Current opinion in

hematology 12 (2): 135–40.doi:10.1097/01.moh.0000153000.09585.79(http://dx.doi.org/10.1097%2F01.moh.0000153000.09585.79) .PMID 15725904 (http://www.ncbi.nlm.nih.gov/pubmed/15725904) .

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^ Laine, R.A. and Rush, J.S. (1988) "Chemistry ofHuman Erythrocyte Polylactosamine Glycopeptides(Erythroglycans) as Related to ABH Blood Groupantigenic Determinants: Evidence that Band 3

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Carbohydrate on Human Erythrocytes Carries theMajority of ABH Blood Group Substance" inMolecular Immunology of Complex Carbohydrates(A. Wu, E. Kabat, Eds.) Plenum PublishingCorporation, N.Y. NY.^ Yazer M, Olsson M, Palcic M (2006). "The cis-ABblood group phenotype: fundamental lessons inglycobiology". Transfus Med Rev 20 (3): 207–17.doi:10.1016/j.tmrv.2006.03.002 (http://dx.doi.org/10.1016%2Fj.tmrv.2006.03.002) . PMID 16787828(http://www.ncbi.nlm.nih.gov/pubmed/16787828) .

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^ Potts, WTW (1979). "History and Blood Groups inthe British Isles". In Sawyer PH. English Medieval

Settlement. St. Martin's Press. ISBN 0-7131-6257-0.

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^ Seltsam A, Hallensleben M, Kollmann A, BlasczykR (2003). "The nature of diversity anddiversification at the ABO locus". Blood 102 (8):3035–42. doi:10.1182/blood-2003-03-0955(http://dx.doi.org/10.1182%2Fblood-2003-03-0955) .PMID 12829588 (http://www.ncbi.nlm.nih.gov/pubmed/12829588) .

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^ Ogasawara K; Bannai M; Saitou N et al. (1996)."Extensive polymorphism of ABO blood group gene:three major lineages of the alleles for the commonABO phenotypes". Human Genetics 97 (6): 777–83.doi:10.1007/BF02346189 (http://dx.doi.org/10.1007%2FBF02346189) . PMID 8641696(http://www.ncbi.nlm.nih.gov/pubmed/8641696) .

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^ a b Calafell, Francesc; et al. (September 2008)."Evolutionary dynamics of the human ABO gene"(http://www.springerlink.com/content/yv4072vu67mv1166/fulltext.html) . Human

Genetics 124 (2): 123–135.doi:10.1007/s00439-008-0530-8 (http://dx.doi.org/10.1007%2Fs00439-008-0530-8) . PMID 18629539(http://www.ncbi.nlm.nih.gov/pubmed/18629539) .http://www.springerlink.com/content/yv4072vu67mv1166/fulltext.html. Retrieved2008-09-24.

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^ CIA World Factbook (https://www.cia.gov/library/publications/the-world-factbook/fields/2119.html)

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^ Blood Types - What Are They?, Australian RedCross (http://www.giveblood.redcross.org.au/page.aspx?IDDataTreeMenu=42&parent=30)

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^ "Austrian Red Cross - Blood Donor Information"(http://old.roteskreuz.at/47_body.html#blutgruppen) .Old.roteskreuz.at. 2006-03-21.http://old.roteskreuz.at/47_body.html#blutgruppen.Retrieved 2010-11-19.

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^ "Rode Kruis Wielsbeke - Blood Donor informationmaterial" (http://www.rodekruiswielsbeke.be/infobloed.html) . Rodekruiswielsbeke.be.http://www.rodekruiswielsbeke.be/infobloed.html.Retrieved 2010-11-19.

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^ Tipos Sanguíneos (http://www.hemoam.org.br29.

/?secao=sangue_tipos)^ Canadian Blood Services - Société canadienne dusang. "Types & Rh System, Canadian BloodServices" (http://www.bloodservices.ca/CentreApps/Internet/UW_V502_MainEngine.nsf/page/Blood%20Types%20and%20Rh%20System?OpenDocument) . http://www.bloodservices.ca/CentreApps/Internet/UW_V502_MainEngine.nsf/page/Blood%20Types%20and%20Rh%20System?OpenDocument. Retrieved 2010-11-19.

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^ Czech Red Cross. "Podíl krevních skupin vpopulaci České republiky" (http://www.cckpraha1.cz/krev.htm) . http://www.cckpraha1.cz/krev.htm.Retrieved 2011-03-18.

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^ Frequency of major blood groups in the Danishpopulation. (http://www.bloddonor.dk/index.php?id=513)

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^ "Veregruppide esinemissagedus Eestis"(http://www.kliinikum.ee/verekeskus/index.php?menu=9&mod=page&id=5) (in Estonian).http://www.kliinikum.ee/verekeskus/index.php?menu=9&mod=page&id=5. Retrieved2010-11-19.

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^ "Suomalaisten veriryhmäjakauma"(http://www.veripalvelu.fi/asp/system/empty.asp?P=1275&VID=default&SID=908029945449597&S=1&C=24395) (inFinnish). 2009-08-21. http://www.veripalvelu.fi/asp/system/empty.asp?P=1275&VID=default&SID=908029945449597&S=1&C=24395. Retrieved2010-11-19.

34.

^ "Les groupes sanguins (système ABO)"(http://www.chpg.mc/go/article.php3?id_article=111)(in French). Centre Hospitalier Princesse GRACE -

Monaco. C.H.P.G. MONACO. 2005.http://www.chpg.mc/go/article.php3?id_article=111.Retrieved 2008-07-15.

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^ de:Blutgruppe#Häufigkeit der Blutgruppen36.^ Blood Donation, Hong Kong Red Cross(http://www5.ha.org.hk/rcbts/doc/neg_newsletter020.pdf)

37.

^ "Blóðflokkar" (http://www4.landspitali.is/lsh_ytri.nsf/pages/bb_blodflokkarnir) (in Icelandic)..landspitali.is. http://www4.landspitali.is/lsh_ytri.nsf/pages/bb_blodflokkarnir. Retrieved 2010-11-19.

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^ "Indian Journal for the Practising Doctor"(http://www.indmedica.com/journals.php?journalid=3&issueid=75&articleid=965&action=article) . Indmedica.com.http://www.indmedica.com/journals.php?journalid=3&issueid=75&articleid=965&action=article. Retrieved 2010-11-19.

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^ "Irish Blood Transfusion Service - Irish BloodGroup Type Frequency Distribution"(http://www.ibts.ie/All_About_Blood

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/Blood_Group_Basics/) . Irish Blood TransfusionService. http://www.ibts.ie/All_About_Blood/Blood_Group_Basics/. Retrieved 2009-11-07.^ "The national rescue service in Israel"(http://www.mdais.org/362/) (in Hebrew). Mdais.org.http://www.mdais.org/362/. Retrieved 2010-11-19.

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^ "Voorraad Erytrocytenconcentraten Bij Sanquin"(http://www.sanquin.nl/Sanquin-nl/erygrafiek.nsf/All/Voorraad-Erytrocytenconcentraten-Bij-Sanquin.html) (in Dutch). http://www.sanquin.nl/Sanquin-nl/erygrafiek.nsf/All/Voorraad-Erytrocytenconcentraten-Bij-Sanquin.html. Retrieved2009-03-27.

42.

^ "What are Blood Groups?"(http://www.nzblood.co.nz/?t=31) . NZ Blood.http://www.nzblood.co.nz/?t=31. Retrieved2010-11-19.

43.

^ Norwegian Blood Donor Organization(https://www.giblod.no/Modules/Page/viewPage.asp?modid=7324&level=7324)

44.

^ "Regionalne Centrum Krwiodawstwa iKrwiolecznictwa we Wroclawiu"(http://rckik.wroclaw.pl/?id=5&go=0) (in Polish).Rckik.wroclaw.pl. 2010-09-02.http://rckik.wroclaw.pl/?id=5&go=0. Retrieved2010-11-19.

45.

^ "Portuguese Blood Institute"(http://www.ipsangue.org/maxcontent-documento-107.html) (in Portuguese). http://www.ipsangue.org/maxcontent-documento-107.html. (assuming Rh andAB antigens are independent)

46.

^ "Frequency of ABO blood groups in the easternregion of Saudi Arabia" (http://cat.inist.fr/?aModele=afficheN&cpsidt=14126617) .Cat.inist.fr. http://cat.inist.fr/?aModele=afficheN&cpsidt=14126617. Retrieved 2010-11-19.

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^ "South African National Blood Service - What'sYour Type?" (http://www.sanbs.org.za/donors_whattype.htm) . Sanbs.org.za.http://www.sanbs.org.za/donors_whattype.htm.Retrieved 2010-11-19.

48.

^ "Federación Nacional de Donantes de Sangre/Lasangre/Grupos" (http://www.donantesdesangre.net/menu.htm) . Donantesdesangre.net.http://www.donantesdesangre.net/menu.htm.Retrieved 2010-11-19.

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^ "Frequency of major blood groups in the Swedishpopulation" (http://www.geblod.nu/general.aspx?PageId=10) . Geblod.nu. 2007-10-02.http://www.geblod.nu/general.aspx?PageId=10.Retrieved 2010-11-19.

50.

^ "Turkey Blood Group Site"(http://www.kangrubu.com/default.asp?sayfa=kan_gruplari) . Kangrubu.com.http://www.kangrubu.com

51.

/default.asp?sayfa=kan_gruplari. Retrieved2010-11-19.^ "Frequency of major blood groups in the UK"(http://www.blood.co.uk/pages/all_about.html) .Blood.co.uk. http://www.blood.co.uk/pages/all_about.html. Retrieved 2010-11-19.

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^ "Blood Types in the U.S"(http://bloodcenter.stanford.edu/about_blood/blood_types.html) . Bloodcenter.stanford.edu.2008-06-20. http://bloodcenter.stanford.edu/about_blood/blood_types.html. Retrieved2010-11-19.

53.

^ RACIAL & ETHNIC DISTRIBUTION of ABOBLOOD TYPES (http://www.bloodbook.com/world-abo.html) , BLOODBOOK.COM

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^ Blood Transfusion Division, United States ArmyMedical Research Laboratory (1971). Selected

contributions to the literature of blood groups and

immunology. 1971 v. 4 (http://books.google.com/?id=ALilcA7Acd0C) . United States Army MedicalResearch Laboratory, Fort Knox, Kentucky.http://books.google.com/?id=ALilcA7Acd0C. "... Innorthern India, in Southern and Central China and

in the neighboring Central Asiatic areas, we find

the highest known frequencies of B. If we leave this

center, the frequency of the B gene decreases

almost everywhere ..."

55.

^ a b Encyclopaedia Britannica (2002). The New

Encyclopaedia Britannica (http://books.google.com/?id=fpdUAAAAMAAJ) . Encyclopaedia Britannica,Inc.. ISBN 0-85229-787-4. http://books.google.com/?id=fpdUAAAAMAAJ. "... The maximum

frequency of the B gene occurs in Central Asia and

northern India. The B gene was probably absent

from American Indians and Australian Aborigines

before racial admixture occurred with the coming of

the white man ..."

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^ Carol R. Ember, Melvin Ember (1973).Anthropology (http://books.google.com/?id=fvpFAAAAMAAJ) . Appleton-Century-Crofts.http://books.google.com/?id=fvpFAAAAMAAJ. "...Blood type B is completely absent in most North and

South American Indians ..."

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^ Laura Dean, MD (2005). Blood Groups an Red

Cell Antigens (http://www.ncbi.nlm.nih.gov/bookshelf/br.fcgi?book=rbcantigen) . NationalCenter for Biotechnology Information, United StatesGovernment. ISBN 1-932811-05-2.http://www.ncbi.nlm.nih.gov/bookshelf/br.fcgi?book=rbcantigen. "... Type A is common in

Central and Eastern Europe. In countries such as

Austria, Denmark, Norway, and Switzerland, about

45-50% of the population have this blood type,

whereas about 40% of Poles and Ukrainians do so.

The highest frequencies are found in small,

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unrelated populations. For example, about 80% of

the Blackfoot Indians of Montana have blood type A

..."^ (PDF) Technical Monograph No. 2: The ABO

Blood Group System and ABO Subgroups

(http://www.biotec.com/pdf/Technical%20Monograph%20No.%202%20-%20ABO%20system%20and%20subgroups.pdf) .Biotec. March 2005. http://www.biotec.com/pdf/Technical%20Monograph%20No.%202%20-%20ABO%20system%20and%20subgroups.pdf. "...The frequency of blood group A is quite high

(25-55%) in Europe, especially in Scandinavia and

parts of central Europe. High group A frequency is

also found in the Aborigines of South Australia (up

to 45%) and in certain American Indian tribes

where the frequency reaches 35% ..."

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^ Sarode, R; Goldstein J, Sussman II, Nagel RL, TsaiHM (June 2000). "Role of A and B blood groupantigens in the expression of adhesive activity of vonWillebrand factor". Br J Haematol. 109 (4): 857–64.doi:10.1046/j.1365-2141.2000.02113.x(http://dx.doi.org/10.1046%2Fj.1365-2141.2000.02113.x) .PMID 10929042 (http://www.ncbi.nlm.nih.gov/pubmed/10929042) .

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^ O'Donnell, J; Laffan MA (August 2001). "Therelationship between ABO histo-blood group, factorVIII and von Willebrand factor". Transfus Med. 11(4): 343–51. doi:10.1046/j.1365-3148.2001.00315.x(http://dx.doi.org/10.1046%2Fj.1365-3148.2001.00315.x) .PMID 11532189 (http://www.ncbi.nlm.nih.gov/pubmed/11532189) .

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^ O'Donnell, J; Boulton FE, Manning RA, Laffan MA(2002-02-01). "Amount of H antigen expressed oncirculating von Willebrand factor is modifiedby ABOblood group genotype and is a major determinant ofplasma von Willebrand factor antigen levels"(http://atvb.ahajournals.org/cgi/content/full/22/2/335). Arterioscler Thromb Vasc Biol. (American HeartAssociation, Inc.) 22 (2): 335–41.doi:10.1161/hq0202.103997 (http://dx.doi.org/10.1161%2Fhq0202.103997) . PMID 11834538(http://www.ncbi.nlm.nih.gov/pubmed/11834538) .http://atvb.ahajournals.org/cgi/content/full/22/2/335.

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^ Gill, JC; Endres-Brooks J, Bauer PJ, Marks WJ,Montgomery RR (June 1987). "The effect of ABOblood group on the diagnosis of von Willebranddisease" (http://www.bloodjournal.org/cgi/content/abstract/69/6/1691) (abstract). Blood 69 (6):1691–5. PMID 3495304(http://www.ncbi.nlm.nih.gov/pubmed/3495304) .http://www.bloodjournal.org/cgi/content/abstract/69/6/1691.

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^ Shima, M; Fujimura Y (1995). "ABO blood groupgenotype and plasma von Willebrand factor innormal individuals". Vox Sang 68 (4): 236–40.doi:10.1111/j.1423-0410.1995.tb02579.x(http://dx.doi.org/10.1111%2Fj.1423-0410.1995.tb02579.x) .PMID 7660643 (http://www.ncbi.nlm.nih.gov/pubmed/7660643) .

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^ Bowen, DJ; Collins PW (March 2005). "Theprevalence of the cysteine1584 variant of vonWillebrand factor is increased in type 1 vonWillebrand disease: co-segregation with increasedsusceptibility to ADAMTS13 proteolysis but notclinical phenotype". Br J Haematol (BlackwellSynergy) 128 (6): 830–6.doi:10.1111/j.1365-2141.2005.05375.x(http://dx.doi.org/10.1111%2Fj.1365-2141.2005.05375.x) .PMID 15755288 (http://www.ncbi.nlm.nih.gov/pubmed/15755288) .

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^ Bongers T; de Maat M; van Goor M et al. (2006)."High von Willebrand factor levels increase the riskof first ischemic stroke: influence of ADAMTS13,inflammation, and genetic variability". Stroke 37(11): 2672–7.doi:10.1161/01.STR.0000244767.39962.f7(http://dx.doi.org/10.1161%2F01.STR.0000244767.39962.f7) .PMID 16990571 (http://www.ncbi.nlm.nih.gov/pubmed/16990571) .

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^ Xie J, Qureshi AA, Li Y, Han J, (2010). ABOBlood Group and Incidence of Skin Cancer(http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0011972). PLoS ONE 5(8): e11972.doi|10.1371/journal.pone.0011972

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^ Wolpin, BM; Kraft, P; Gross, M; Helzlsouer, K;Bueno-De-Mesquita, HB; Steplowski, E;Stolzenberg-Solomon, RZ; Arslan, AA et al. (2010)."Pancreatic cancer risk and ABO blood groupalleles: results from the pancreatic cancer cohortconsortium" (http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=2943735) .Cancer research 70 (3): 1015–23.doi:10.1158/0008-5472.CAN-09-2993(http://dx.doi.org/10.1158%2F0008-5472.CAN-09-2993) . PMC 2943735(http://www.pubmedcentral.gov/articlerender.fcgi?tool=pmcentrez&artid=2943735) .PMID 20103627 (http://www.ncbi.nlm.nih.gov/pubmed/20103627) .http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=2943735.

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^ Amundadottir, L; Kraft, P; Stolzenberg-Solomon,RZ; Fuchs, CS; Petersen, GM; Arslan, AA; Bueno-

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De-Mesquita, HB; Gross, M et al. (2009)."Genome-wide association study identifies variantsin the ABO locus associated with susceptibility topancreatic cancer"(http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=2839871) .Nature genetics 41 (9): 986–90. doi:10.1038/ng.429(http://dx.doi.org/10.1038%2Fng.429) .PMC 2839871 (http://www.pubmedcentral.gov/articlerender.fcgi?tool=pmcentrez&artid=2839871) .PMID 19648918 (http://www.ncbi.nlm.nih.gov/pubmed/19648918) .http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=2839871.^ Gates, MA; Wolpin, BM; Cramer, DW; Hankinson,SE; Tworoger, SS (2010). "ABO blood group andincidence of epithelial ovarian cancer"(http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=2946962) .International journal of cancer. Journal

international du cancer 128 (2): 482–486.doi:10.1002/ijc.25339 (http://dx.doi.org/10.1002%2Fijc.25339) . PMC 2946962(http://www.pubmedcentral.gov/articlerender.fcgi?tool=pmcentrez&artid=2946962) .PMID 20309936 (http://www.ncbi.nlm.nih.gov/pubmed/20309936) .http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=2946962.

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^ a b Blood Group A Suptypes72.

(http://web.archive.org/web/20080802232301/http://www.owenfoundation.com/Health_Science/Blood_Group_A_Subtypes.html) , The OwenFoundation. Retrieved 2008-07-01.^ Erb IH (1 May 1940). "Blood GroupClassifications, a Plea for Uniformity"(http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=537907) .Canadian Medical Association Journal 42 (5):418–21. PMC 537907(http://www.pubmedcentral.gov/articlerender.fcgi?tool=pmcentrez&artid=537907) .PMID 20321693 (http://www.ncbi.nlm.nih.gov/pubmed/20321693) .http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=537907.

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^ Liu, QP; Sulzenbacher G, Yuan H, Bennett EP,Pietz G, Saunders K, Spence J, Nudelman E, LeverySB, White T, Neveu JM, Lane WS, Bourne Y,Olsson ML, Henrissat B, Clausen H (April 2007)."Bacterial glycosidases for the production ofuniversal red blood cells" (http://www.nature.com/nbt/journal/v25/n4/abs/nbt1298.html) . Nat

Biotechnol 25 (4): 454–64. doi:10.1038/nbt1298(http://dx.doi.org/10.1038%2Fnbt1298) .PMID 17401360 (http://www.ncbi.nlm.nih.gov/pubmed/17401360) . http://www.nature.com/nbt/journal/v25/n4/abs/nbt1298.html.

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^ American Red Cross, Southern California BloodServices Region (n.d.). "Answers to CommonlyAsked Questions About Blood and Blood Banking"(http://www.socalredcross.org/pdf/BloodThe.pdf)(PDF). Blood: the Basics: 4.http://www.socalredcross.org/pdf/BloodThe.pdf.Retrieved 2007-11-16.

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^ http://www.dadamo.com/76.^ Klein, Harvey G (March 7, 2005). "Why DoPeople Have Different Blood Types?"(http://www.sciam.com/article.cfm?id=why-do-people-have-differ) . Scientific American.http://www.sciam.com/article.cfm?id=why-do-people-have-differ. Retrieved 2007-11-16.

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Further reading

Dean L (2005). "Chapter 5: The ABO blood group." (http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=rbcantigen.chapter.ch05ABO) . Blood Groups and Red Cell Antigens.http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=rbcantigen.chapter.ch05ABO. Retrieved 2007-03-24.Farr A (1 April 1979). "Blood group serology--the first four decades (1900--1939)"(http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1082436) . Med Hist

23 (2): 215–26. PMC 1082436 (http://www.pubmedcentral.gov/articlerender.fcgi?tool=pmcentrez&artid=1082436) . PMID 381816 (http://www.ncbi.nlm.nih.gov/pubmed/381816) .


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http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1082436.

External links

ABO (http://www.ncbi.nlm.nih.gov/gv/mhc/xslcgi.cgi?cmd=bgmut/systems_info&system=abo) atBGMUT Blood Group Antigen Gene Mutation Database at NCBI, NIHEncyclopaedia Britannica, ABO blood group system (http://www.britannica.com/eb/article-9003372/ABO-blood-group-system)National Blood Transfusion Service (http://www.blood.co.uk/pages/world_blood.html)Molecular Genetic Basis of ABO (http://abobloodgroup.googlepages.com)

Retrieved from "http://en.wikipedia.org/wiki/ABO_blood_group_system"Categories: Blood antigen systems

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