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Journal of Feline Medicine and Surgery(2002)4, 69\u201376
doi:10.1053/jfms.2001.0162, available online at http://www.idealibrary.com on
REVIEW
The feline AB blood group system and its importance
in transfusion medicine
CM Knottenbelt

Lecturer in Internal Medicine,
Department of Veterinary Clinical
Studies, University of Glasgow
Veterinary School, Bearsden Road,
Bearsden, Glasgow G61 1QH, UK

Date accepted: 16 November 2001

he feline blood group system was \ufb01rst characterised in 1962 following the identi- \ufb01cation of two major blood types (desig-

nated A and B) (Eyquem et al 1962). One blood

group system with three types (A, B and AB) has since been identi\ufb01ed (Auer & Bell 1981). Naturally occurring antibodies against foreign blood types in the cat are responsible for the premature destruction of transfused red cells, clinically severe transfusion reactions and neonatal isoerythrolysis.

Type A cats given type B blood may develop a mild transfusion reaction that is often not clini- cally apparent. The recipient\u2019s packed cell vol- ume (PCV) falls to pre-transfusion levels within days of the transfusion (Giger & Bucheler 1991). Type B cats that are transfused with type A blood invariably develop rapid, potentially fatal trans- fusion reactions following even a single transfu- sion of a small volume of blood (Auer & Bell

1981).

This paper reviews current understanding of feline blood groups and alloantibodies, and their importance in transfusion medicine.

Feline blood group prevalence

In world-wide studies, type A has consistently been found to be the most common blood type. The proportions of type B cats, however, have shown considerable geographical variations

(Eyquem et al 1962,Auer & Bell 1981,Ejima

et al 1986, Giger et al 1989, Giger et al 1991a, 1991b, Continenza et al 1992, Giger et al 1992, Knottenbelt et al 1999a) (Table 1). The frequency

of type B cats varies dramatically in different breeds (Table 2), however type AB cats are con- sistently rare (Auer &Bell 1981,Giger et al 1991a,

1991b, Griot-Wenk et al 1996, Knottenbelt et al
1999a). Wild felids have recently been reported
to have the same blood groups as domesticated
cats (Griot-Wenk & Giger 1999).
Blood group inheritance
Feline blood types are determined by at least two
alleles (a or b) at the same gene locus (Giger et al
1991a). The type a allele appears to be com-

pletely dominant over the type b allele. Therefore cats with phenotype A may have the genotype a/a or a/b, whilst only cats homozygous for the B allele express signi\ufb01cant amounts of the type B erythrocyte antigen (Giger et al 1991a) and none of the type A antigen. With the exception of dogs and cats, inheritance of blood types within a blood group system in most other species is typically by codominant alleles (Symons & Bell

1992). Cats are therefore unusual in inheriting
type B as a recessive trait (Rippee et al 1989,
Giger et al 1991a).
The AB blood type was previously thought to
result from matings between two phenotypic
1098-612X/02/020069+08 $35.00/0
\u00a9 2002 ESFM and AAFP. Published by Elsevier Science Ltd. All rights reserved.

type A cats both with the genotype a/b, or between a phenotypic type A cat (genotype a/b) and a type B cat, or between two type B cats (Griot-Wenk & Giger 1991). However, more

recent work suggests that type AB is inde- pendently inherited in the feline blood group system, since type AB cats are rarely the off- spring of matings between type A and type B

Table 1.Frequency of blood types in domestic short and long-haired cats world-wide
Country
n
Type A
(%)
Type B
(%)
Type AB
(%)
References
Australia
1895
73.3
26.3
0.4
Auer & Bell (1981)
Austria
101
97
3
0
Giger et al (1992)
England
477
97.1
2.9
0
Holmes (1950)
Finland
61
100
0
0
Giger et al (1992)
France
350
85.1
14.9
0
Eyquem et al (1962)
Germany
600
94.0
6.0
0
Giger et al (1992)
Holland
95
95.8
3.1
1.1
Giger et al (1992)
Italy
401
88.8
11.2
0
Giger et al (1992)
Japan
265
89.3
1.0
9.7
Ejima et al (1986)
Scotland
70
97.1
2.9
0
Giger et al (1992)
137
87.6
8.0
4.4
Knottenbelt et al (1999a)
Switzerland
1018
99.6
0.4
0
Giger et al (1992)
USA
1072
99.7
0.3
0
Giger et al (1989)
3785
98.1
1.7
0.1
Giger et al (1991b)
Note: n=number of cats tested; %=percentage of cats.
Table 2.Frequency of feline blood types in different breeds worldwide
Breed
Country
n
Type A
(%)
Type B
(%)
Type AB
(%)
Abyssinian
USA
194
79.9
20.1
American short-hair
USA
15
100.0
Bengal
UK
8
50.0
50.0
Birman
USA
216
82.4
17.6
UK
11
72.7
9.1
18.2
British short hair
USA
85
41.2
58.8
UK
105
41.0
57.1
1.9
Burmese
USA
25
100.0
UK
6
83.3
16.7
Devon Rex
USA
100
57.0
43.0
UK
2
100.0
Norwegian Forest
USA
20
100.0
Oriental short hair
USA
15
100.0
Persian
USA
170
75.9
24.1
UK
16
87.5
12.5
Italy*
38
97.4
2.6
Germany\u2020
25
84.0
16.0
Ragdoll
UK
2
100.0
Scottish Fold
USA
27
85.2
14.8
Siamese
USA
99
100.0
UK
4
100.0
Somali
USA
27
77.8
22.2
UK
9
77.8
22.2
Tonkinese
USA
31
100.0
Note: n=number of cats tested; %=percentage of cats.
Figures taken fromGiger et al 1991a (USA) andKnottenbelt et al 1999a (UK) except:
*Continenza et al 1992; \u2020Von Haarer & Grunbaum 1990.
70
CM Knottenbelt
cats (Griot-Wenk et al 1996). The ab allele

appears to be recessive to the a allele but domi- nant over the b allele (Griot-Wenk et al 1996). Type AB cats are only found in breeds in which type B cats have been identi\ufb01ed (Griot-Wenk

et al 1996).
Blood group antigens

The antigens in the feline AB blood group are not serologically related to human ABO blood group antigens (Auer & Bell 1981). By 38 days of gestation, type A and type B antigens are present on foetal erythrocytes (Auer & Bell 1981). Blood type antigens are thought to be determined by the form of neuraminic acid attached to the glycolipids of the red cell (speci\ufb01cally the ganglioside GD3), and possibly also a glycopro- tein of the erythrocyte membrane (Andrews

et al 1992). Type A blood has NeuGc-NeuGc-
Galactose-Glucose-Ceramide ([NeuGc]2

GD3) (where NeuGc represents N-glycolylneuraminic acid) as the major glycolipid (Butler et al 1991a). N-acetylneuraminic acid (NeuAc) is, however, also present in variable proportions in type A blood. This variability may be due to the homozygous or heterozygous nature of cats with type A blood (Griot-Wenk et al 1993). Type B blood, in contrast to type A blood, expresses no NeuGc, the major glycolipid being NeuAc- NeuAc-Galactose-Glucose-Ceramide ([NeuAc]2 GD3) (Butler et al 1991a). Type B erythrocytes are therefore characterised by [NeuAc]2GD3, the presence of NeuAc on the 50 kD erythrocyte membrane glycoprotein and a lack of detectable NeuGc (Andrews et al 1992).

Type AB blood shows co-expression of both
type A and type B antigens (Andrews et al 1992,
Griot-Wenk et al 1996). The proportions of

NeuAc and NeuGc, however, vary between dif- ferent type AB individuals (Andrews et al 1992) and intermediate forms of glycolipids have been recovered from type AB red cells (Griot-Wenk

et al 1996). Cats without either type A or B red
cell antigens (analogous to the human blood
group O) have not been reported.

It has been suggested that the enzyme NeuAc hydroxylase is responsible for the conversion of NeuAc to NeuGc in type A cats, and that type B cats lack this enzyme (Butler et al 1991b,

Andrews et al 1992). Type AB cats may have

a hereditary mutation of the enzyme or a gene that alters enzyme expression. This theory is supported by the \ufb01nding that type AB cats are only produced from crosses involving an AB

phenotype (Auer & Bell 1981). This theory how- ever, fails to account for the \ufb01nding of variable amounts of NeuAc in type A cats (Griot-Wenk

et al 1993), unless enzyme activity was limited by

an alternative mechanism in some type A cats. Measurement of NeuAc hydroxylase activity in cats has not been performed.

Alloantibodies

The proportion of type A cats with low levels of naturally occurring antibody directed against type B red cells varies geographically (Auer &

Bell 1981, Ejima et al 1986, Giger et al 1989,
Bucheler & Giger 1993, Knottenbelt et al 1999b).

However all type B cats appear to have high levels of naturally occurring anti-A antibody (Giger et al 1989,Knottenbelt et al 1999b). Cats with type AB blood do not have alloantibodies (Griot-Wenk et al 1996). The naturally-occurring anti-B antibodies of type A cats are weak IgM agglutinins and weak haemolysins comprising equal proportions of IgM and IgG (Bucheler &

Giger 1993). Anti-A alloantibodies appear to be
strong anti-A haemolysins and haemagglutinins
predominantly of the IgM class (Wilkerson et al
1991, Bucheler and Giger 1993), although some
haemagglutinating activity has also been
associated with IgG (Wilkerson et al 1991).

Naturally-occurring alloantibodies are be- lievedto result from exposure to epitopes that are commonly found in nature (usually as structural components of a variety of organisms including plants, bacteria and protozoa), and that are simi- lar or identical to blood group antigens (Male

1996, Tizard 1996). In man, the epitopes respon-
sible are thought to be microbial antigens present
on intestinal bacteria (Male 1996,Kuby 1997,
1997b). In cats however, the origin of these

epitopes has not been established. Exposure to these epitopes results in the formation of anti- bodies against antigens that the individual does not naturally possess. The antibody produced will then cross-react with similar antigens (such as blood group antigens) found on foreign eryth- rocytes (Kuby 1997a). Exposure to epitopes that are similar to self antigens will not result in antibody formation due to self tolerance (Kuby

1997b). This mechanism explains the reported
absence of alloantibodies in type AB cats (Auer &
Bell 1981), since the antigens found in nature that

resemble either type A or type B blood group antigens will be recognised as \u2018self\u2019. The absence of naturally-occurring alloantibodies in many domestic animal species may be due to an

Blood groups and their importance in transfusion
71
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