452 
HEMATOLOGY 
Experi- 
ment 
Serum 
T- ouabain 
(Ml/ 
preincubation 
K*-pump flux 
(mM/(L cells) 
X hr] 
Inhibition 
(%l 
Molecules 
T-ouabain/ 
cell 
Pump 
sites/ 
cell 
E 84 
Anti ■ L 
0 
0.28 
10-' 
0.12 
57 
70 
122 
HK Nl 
0 
0.11 
10-' 
0.065 
41 
40 
97 
E 86 
Anti - L 
0 
0.26 
10-' 
0.076 
71 
45 
63 
HK Nil 
0 
0.001 
10-' 
<0 
71 
21 
30- 
E88 
Anti L 
0 
0.38 
10-' 
0.25 
37 
30 
82 
HK Nl 
0 
0.094 
10-' 
0.059 
37 
10 
27 
* Since no K'*'-pump flux was measured in this part of exp. 86, the degree of inhibition by 10- 'M 
T-ouabain could not be estimated. Therefore, the number of pump sites was computed on the basis of 
71% K'*'-pump flux inhibition found in presence of anti- L. 
Figure 3. — Effect of L-antiserum on the number of 
ouabain molecules bound per LK sheep red celP' 
terized the anti-Lantibody as an IgG immuno- 
globulin. The same investigators have shown 
that the F(ab)o antibody fragment alone is ca- 
pable of stimulating potassium influx in LK 
cells. The physical and chemical characteristics 
of the M and L antigens are still unknown. 
CATTLE 
At least 11 major blood grouping systems 
have been identified in cattle: A, B, C, F-V, J, 
L, M, N, S, Z, and - (Figure 4). Some 
laboratories report more major bovine blood 
groups than those listed here. However, with 
the exception of N, these are the most widely 
recognized. All bovine blood typing has been 
done with lytic techniques since bovine erythro- 
cytes tend not to agglutinate even when they 
have been incubated with multiple sensitizing 
doses of blood typing antibodies. Rabbit and 
guinea pig complement are used in routine typ- 
Genetic 
systems 
Year 
discovered 
Number of 
blood factors 
Number of 
phenogroups 
Number of 
distinguishable 
phenotypes 
A 
1944 
5 
10 
13 
B 
1940 
50 
300 + 
15,000? 
C 
1941 
10 
35 + 
200? 
F-V 
1943 
5 
4 
9 
J 
1942 
2 
4 
4 
L 
1947 
1 
2 
2 
M 
1958 
2 
3 
3 
N 
1960 
1 
2 
2 
S 
1943 
6 
5 
12 
z 
1941 
1 
2 
3 
R'-S' 
1960 
2 
2 
3 
Figure 4. — Major blood groups in cattle' 
ing of bovine blood. At present over 100 blood 
typing reagents are used in analyzing cattle 
blood groups. 
The B system was the first bovine blood 
group to be extensively studied. It has be- 
come one of the most complex blood groups 
known in any species. Group B was first recog- 
nized as a multiple allelic system in 1940 by 
Stormont who observed that bovine blood factor 
K appeared only in the presence of factors B 
and G. He proposed that the observed pheno- 
types BGK, BG, B, G, and no-BGK were con- 
trolled by multiple alleles which he designated 
IjBGK^ Ijbg^ Ijb^ 1)6^ and b~. Here the superscripts 
BGK, BG, B, G, and " are referred to as 
phenogroups of the B system. It soon became 
apparent that many other blood factors were in 
fact part of the B system^o-^s including such 
phenogroups as BGKOoYiA'E',-, G"K'Y'B'0', 
OiTiE'.^F', K', PQE'i, GO.b', and BGKOxYiA'. 
During the past 30 years more than 300 pheno- 
groups have been recognized in the B system. 
There are more than 45,000 possible diploid com- 
binations of B phenogroups.^^ How many of 
these would produce serologically distinguish- 
able phenotypes is anybody's guess. The range of 
possibilities continues to grow as new alleles are 
discovered, some probably appearing as a result 
of spontaneous mutation. 
The J antigen was discovered in 1942 by Fer- 
guson, Stormont, and Irwin^i working with 
naturally occurring antibodies from the blood 
of what were later to be classified as J negative 
cattle. Soon thereafter a soluble substance was 
found in the serum of J positive cattle which in- 
hibited the reaction of cellular J substance with 
anti-J antisera. On the basis of these observa- 
tions, Stormont proposed that J substance is pri- 
marily a soluble antigenic entity elaborated in J 
positive cattle by cells outside the circulatory 
system. J substance has been found in areas 
outside of the circulation including the saliva, 
gastric mucosa, and urine. It is adsorbed by the 
erythrocytes from the serum. The ability of cat- 
tle erythrocytes to adsorb J substance was dem- 
onstrated when erythrocytes from J negative 
individuals were incubated in J positive serum. 
In the presence of the J positive serum, J nega- 
tive cells became serologically J positive. Stone 
and Irwin-'^ later discovered cattle with eryth- 
