THE JU BLOOD TYPING SYSTEM OF THE SPERM WHALE
AND SPECIFIC SOLUBLE SUBSTANCESJOHN E. CUSHING,* KAZUO FUJINOAND NORA CALAPRICE*
The use of immunogenetic concepts in marine population research is rapidly expanding (Cushing 1962) and pertinent material on whales has been reviewed (Fujino,1960). In the case of sperm whales, a "species-specific" antigen resembling thehuman B antigen has been found on their erythrocytes, and antibodies for thehuman ABO system occur variably in their serums. Chicken and rabbit antiserums
prepared against sperm whale erythrocytes have been used to distinguish six
phenotypes among twenty-six individuals. One of the antigens designated, Sp,
could also be recognized by a natural antibody in sei whale serum (Fujino 1954).
The present paper reports additional work on sperm whale antigens that wasdone in California (Cushing and Calaprice), in the Antarctic (Fujino) and in Peru
(Cushing).
CALIFORNIA RESEARCH

Samples of bloods from ten sperm whales taken by San Francisco shore whaling
stations were preserved by glycerol-freezing (Cushing et al., 1959). Erythrocytes
were recovered by dialysis and compared with those of finback, humpback, andsei whale erythrocytes with respect to their reactions with Fujino's (1956) heteroimmune anti-finback Jul (no. 47) and Ju2 (no. 34) serums. These comparisonsdemonstrated that, as in the humpback, pygmy-blue and sei whales, antigens occurin sperm whales that are very similar in specificity to the Jul and Ju2 finback
antigens. These antigens behave in finbacks as if determined by a pair of allelicgenes (Fujino, 1960).
The normal serums of horses, cattle, sheep and pigs were found to be usefulreagents for detecting the Ju2 antigen. Antibodies for this antigen occurring in allserums examined. The serum of the California spiny lobster (Panulirus interuptus)
was found to contain antibodies capable of agglutinating some Ju2-positive cells,
but not others, suggesting that it may prove to be valuable as a Ju2 subtyping
reagent (These reactions will be reported in a separate paper).
Isoantibodies in Ju2-negative sperm whales for Ju2-positive erythrocytes
were found, as were cross-reactions between the Ju2-positive cells of one species
with the isoantibodies of other species. These results suggest that the Sp antigen
already referred to may have been Ju2 because of its agglutinability with sei whale
serum. No other relations between the sperm whale antigens first described by
Fujino and the ones described in this paper as like those of the finback Ju system
can be determined, as the earlier and-sperm whale reagents are no longer available* Department- of Biological Sciences, The :University of California, Santa Barbara, California, U.S.A.
67

J. E. CUSHING, K. FUJINO AND N. CALAPRICE

68

for comparison.
The results given in the above section, together with relevant observations
reported in earlier papers, establish the possibility of using the readily available
reagents in normal serums for comparison of Ju2-positive reactions among spermand other whale species in different parts of the world. The results of two suchstudies are given below.
ANTARCTIC RESEARCH

Data on the Ju system of blood types in Antarctic populations of sperm whales was
obtained by Fujino while he served as biologist aboard the factory ship, Kyokuyomaru No. 2 during the 1960/61 whaling season. Samples were taken from theheart of all but a few whales at an average time of eight hours post-mortem in sea
and air temperatures which fluctuated within a few degrees of zero centigrade.
The exceptional samples came from large clots of blood within the body cavity.
Nine volumes of sample were added to one volume of the usual collecting solution.
This consistsof8.5 gm. NaCl, 50 gm. of sodium citrate and 0.5 gm. of the antimicrobial
TABLE I.

Undiluted
serums:ISOAGGLUTINATION OF VARIOUS WHALE ERYTHROCYTESBY UNDILUTED SERUMS*
Ju2-negativeJu2-positive cells
cells,----''----.

Ju2-neg.
serum{ 15495

Ju2-pos.
serum

163
152
IOI
110
124
111
120
100

l

100

6

8

120

124

110

IOI

Ill

90

163

7

-ttt
-ttt

-ttt

-ttt

-ttt
-ttt

-ttt
-ttt

-ttt
-ttt

-ttt
-ttt

-ttt
-ttt

+
*
±
* * *

±

*
** *+ **+ *-ttt *+ ** *++ + -ttt -ttt
+ + +
* * ±

-ttt

78

-ttt

152

95

154

* See absorptions in Table 2.
These reactions show that subtypes exist among Ju2-positive cells and that the weakersubtypes can contain isoantibodies for stronger ones. Serums and cells of whales 95 and 154
are representative of those of all Ju 2 negative whales -tit symbolizes strong agglutination, ranging
throughand + to - for no agglutination.

*

guanofuracin per liter of water. These samples remained in good condition for
thirty days or longer under refrigeration. Serums of such samples showed little
hemolysis, and had no opportunity to become mixed with other tissue fluids.
Each sample was tested with the same normal bovine serum, using conventional
saline agglutination techniques. Agglutination was taken as evidence of thepresence of Ju2 antigen. All Ju2 positive samples, together with various negative
ones, were subsequently compared as to their reactions with normal horse serum,
the same unabsorbed anti-finback serums used by Cushing and Calaprice, and theJU BLOOD TYPING SYSTEM AND SOLUBLE SUBSTANCES

69

TABLE 2. EXAMPLES OF ABSORPTIONS OF CATTLE SERUM(NO. 25) THAT SHOW SUBTYPE DIFFERENCES AMONG
JU 2-POSITIVE SPERM WHALES*
Reactions with absorbed serums diluted
one in eight
Absorbing
Test Cells
Cells:
100
124
111
7
95

Reactions with unabsorbed serum, dilutions:
Cells:
1: 2 1: 4
1: 8 1 : 16 1 : 32 1: 64

Ju2
pos.

('00

124
111
7

Ju2
neg. { 95

tit
tit
tit
tit

tit
tit
tit
+t-

tit
tit
tit

tit
tit
+t-

+t+t-

+

+
+

r
124
111
7

+ttit
tit

tit
tit

tit

Ju2 { 95
neg.

tit

tit

tit

Ju2
pos.

+

* Also see Table 1 for reactions of these red cells with various whale normal serums.

70

Is.

KERGUELEN Is.

Fig. 1. Localities where sperm whales were taken by Kyokuyo-maru No. 2. in the
1960/61 whaling season.

+

]. E. CUSHING, K. FUJINO AND N. CALAPRICE

70

isoantibodies in a single sperm whale serum. These comparisons continued toconfirm the usefulness of normal animal serums as reagents for the detection of theJu2 antigen on erythrocytes.
Isoantibodies for Ju2-positive cells were found in all Ju2-negative serums examined (one hundred and twenty-seven). In the weaker Ju2-positive subtypes
were detected isoantibodies for the stronger Ju2 subtypes as showin in Table 1.
Isoantibodies for Jul-positive cells were not found.
All Ju2-positive samples from areas A and B (see Fig. 1), having been collected within a span of twenty-eight days, were compared with respect to theirability to absorb bovine serum. These absorptions showed that Ju2-positive cells
exist as a series of at least four subtypes (Table 2). No subtypes of Ju2-negative
cells were found following absorption of antifinback Jul reagent with cells fromforty-six negative samples. Experiments were performed on the reactions ofantibodies eluted from erythrocytes following absorption of bovine and anti-finback
Jul serum. These will be reported in detail in connection with other related
studies.
One whale was found that appeared to be a blood group chimera in that its
cells consisted of a mixture of Ju2-positive and Ju2-negative cells as determined by
microscopic examination of agglutionations, and by hemolytic tests.
TABLE 3.

FREQUENCY OF OCCURRENCE OF BLOOD TYPES OF THE
ANTARCTIC SPERM WHALES*
Area A

Blood types
Ju2-negatives

Area B

Area CSum of all whales

~

~~

~

Act. no.

Freq.

Act. no.

Freq.

Act. no.

Freq.

Act. no.

Freq.

105

0.898

95

0.969

15

0.882

215

0.927

:

) 0 031

98

I.OOO

Ju2 1

Ju2-positives,
separated bysubtypes.**

l
J

Ju22

2

J u2

8

0.102
3

0.073

Ju2 4
Sum

117

I.OOO

17

I.OOO

232

I.OOO

* See Fig. I for area limits* * Symbols of four subtypes of Ju2-positives were named in accordance with those in finbacks(Fujino, 1962).

The frequencies of Ju2-positive cells in population samples from the different
Antarctic areas shown in Fig. 1 are given in Table 3. These are arranged
accrding to subtype differences and suggest that larger samples will show that the
sperm whale population of area A may differ from those of Band C.
PERUVIAN STUDIES

Sperm whale Ju blood types were studied in Peru by Cushing during July-August
1960 at the shore whaling station of Cia Balleneria Del Norte, Tierra Colorado

JU BLOOD TYPING SYSTEM AND SOLUBLE SUBSTANCES

71

Bay, Paita, Peru. Whales were fished twenty to ninety miles off-shore and usually
had been dead twelve to thirty hours before flensing. Blood samples were takenfrom various flensing cuts in the musculature, it being impractical to obtain heart
blood as was done by Fujino. Whales in which decomposition was apparent were
not sampled.
The erythrocytes obtained were in good condition, but the serum showed much
hemolysis and was subject to mixt with fluids from the muscle tissue. A few milligrams of streptomycin and sulfathiazole were mixed with each 100 ml. sample,
which kept well for several days refrigerated.
Conventional slide agglutination tests were used, being scored in a range between - for no reaction through itlt for the strongest agglutination. (Note that thescore of agglutinations given by Fujino does not extend beyond tit. Study of thesame materials by both authors while together in Tokyo showed this difference inconvention had no effect on interpretations of the data obtained). Reagents used
included the same anti-finback and stock of normal domestic animal serums referred to in the foregoing sections of this report. Emphasis on the Peruvian work
centers on the reactions of erythrocytes with a pool of horse serum. As in previous
work, saline controls were run on all cell suspensions to avoid possible error throughautoagglu tionation.
The horse serum readily distinguished two types of individuals. Cells reacting
strongly were classified as Ju2-positive, those at dilutions of serum at one in five or
less as Ju2-negative. Reactions with the anti-fin whale heteroimmune serums
confirmed this classification.
As the first whale typed gave a strong positive reaction and the next three
gave negative reactions, it looked as if the frequency of Ju2-positive cells would
be readily obtained. However, a new phenomenon was encountered early in theinvestigation that showed this would be impossible to achieve within the time
available. This phenomenon was the observation that five different samples which
initially reacted as Ju 2-negative, changed type over a period of days and became
Ju2-positive. In contrast, eight other Ju2-negative cells did not change type,nor did any Ju2-positive cells become negative.
As the changes occurred in spite of refrigeration and the presence of antibiotics, were Ju specific, and took place in samples in good condition, it seemed unlikely that they were due to microbial actions (Stormont et al., 1960).
The possibility that the erythrocytes had acquired a specific soluble Ju2
substance (hereafter Ju2 SSS) from the serum or tissues was therefore investigated.
Such substances were first described by Stormont who showed that the J types ofcattle erythrocytes are determined by SSS in the blood plasma (Stormont, 1949).
They have subsequently been found in man, sheep and other domestic species (Anintroduction to recent literature on soluble blood group antigens will be found in
Sneath et al., 1959, Stone, 1962, Stormont et al, 1960, and Race et al., 1958).
Limited observations on sperm whale human B-like antigen and on the finback Ju
system have already suggested their occurrence in cetaceans (Cushing et al., 1959,
~~l~.
.
.

]. E. CUSHING, K. FUJINO AND N. CALAPRICE

72

Table 4 shows the reactions of the whales that changed type. While circumstances did not permit an extensive series of experiments, several kinds of observations were made which indicate that the type of change was actually due to postmortem coating of erythrocytes with Ju2 SSS.
The first of these observations was the demonstration that Ju2 SSS capableof inhibiting the reactions of horse serum withJu2-positive cells occurs in the serumsof whales that changed types, and in examples of other positive whales (Table 5).
The failure of inhibition with respect to certain combinations of cells and serums
can be reasonably explained as due to subtype differences among the reactants
involved. Serums of four whales typed as Ju2-negative did not inhibit the action
TABLE 4.
Whale
nos:

3
3
*3
19
19
19
21
21
21
21
5-5
5-5
5-5
5-5
4-3
4-3

REACTIONS SHOWING CHANGE IN JU TYPE OF THE
ERYTHROCYTES OF INDIVIDUAL WHALES
Date tested,

Horse serum dilutions

1:5

1: 10

1: 20

-fttt

-fttt

1: 160

1: 40

1: 80

+H-

*

+

-fttt
-fttt

-fttt
-fttt

-fttt
-fttt

+H-fttt*+****-fttt

+H-

*

+

+

±

*-fttt

*+H-

+
-fttt
-fttt

*
+
*+H-

-fttt

-fttt

-fttt

-fttt

+

+

*

1960
7-31
8-2
8-18
8-3
8-5
8-15
8-3
8-5
8-7
8-15
8-6
8-7
8-9
8-13
8-5
8-9

+
* This sample represents cells recovered from an aliquot of 3 blood frozen in glycerol citrate
-fttt

-fttt

+H-

+H-

and recovered by dialysis Aug. 18. The cells had not become positive during this time, incontrast to the change in unfrozen cells.

of horse serum. Account was taken of the occurrence and action of isoantibodies
in these and other serums where these antibodies might have complicated results.
Another kind of supporting evidence are the results of some absorptions ofhorse serum involving the erythrocytes of some of the whales that changed types,
and which are shown in Table 5 to have J u2 SSS in their serums (Table 6). Theseabsorptions show that, as in the inhibition tests, individual variations exist in thereactions of different erythrocytes with different serums. The relations between
19 and 21 are strikingly similar in both kinds of experiments. Cells 4-1 and 5-6
removed antibodies agglutinating some other kinds of cells while not agglutinating
to any degree in unabsorbed horse serum themselves ; also that the serums of thesecells were capable of inhibition (Table 5). This is a situation similar to that reported
for certain of the cattle J types (Stormont et al., 1960, Stormont, 1962).
A third kind of experiment further supports the possibility that the type changes

JU BLOOD TYPING SYSTEM AND SOLUBLE SUBSTANCES

73

TABLE 5. EXAMPLES OF THE INHIBITION OF HORSE SERUM BY
SERUMS FROM JU2-POSITIVE WHALES. THOSE THAT CHANGEDTYPE ARE INDICATED BY AN ASTERISK
Test
cells

horse
serum
only

5-5
4-3
21
0-1
19

1tH1tH1tH1tH1tH-*21

1tH1tH-

*4-3

*5-5

*0-3

0-1

5-6

4-1

*19

+
+

+
+

+
+

+
+

*

1tH1tH1tH-

-Ht

1tH-

**
*
*

*
*
*
*

1tH1tH1tH-

1tH1tH-

1tH-

+
+

1tH-

+

*

Horse serum diluted one in ten plus individual whale serums (excepting first column whichshows the reactions of horse serum alone). One drop of whale serum was added to one drop ofhorse serum for each test. Cells were added fifteen minutes later and readings made after an
additional twenty minutes. -Ht indicates strong agglutination and therefore no inhibition, while-indicates no agglutination and therefore complete inhibition. Note that homologous combinations
4-3, 5-5 and 19 did not show complete inhibition. No isoagglutinins for the cells used in theabove tests were found in serums 21, 5-5, 0-1, 5-6, and 19. Serums 0-3, 4-3, and 4-5 were
not tested for isoagglutinins. Four Ju2-negative serums (0-2, 0-4, 4-5, and 11-7) were found tocontain isoagglutinins for the above cells, and did not inhibit the agglutination of these cells byhorse serum.
TABLE 6.

THE REACTIONS BETWEEN VARIOUS ABSORBEDHORSE SERUMS AND ERYTHROCYTES

Absorbing
cells

5-6

4-5

4-1

19

21

Test cells
4-4
4-3

Unabsorbed
serum (1 : 10)

none

+

1tH-

1tH-

1tH-

±

+

1tH-

1tH-

Absorbed
serums (1 : 10)

19 Ju2-pos.
21 Ju2-pos.
4-1 Ju2-neg.
5-6 Ju2-neg.
4-5 Ju2-neg.

1tH-*
1tH1tH--Ht+

1tH1tH-

*

8-3

4-2

5-5

6-2

1tH-

1tH-

1tH-

1tH-

-Ht

1tH-

1tH-

1tH-

1tH-

+

-Ht

-Ht

* * *

1tH-

Blank spaces in columns to right show that no tests were made. Serums were absorbed at room
temperature with one-third volume of packed cells for thirty minutes. Results show the marked
reciprocal relationship between 19 and 21 observed in inhibitions (table 5). Note that absorption
with cells of 19 was not complete. Other heterogeneities are revealed suggestive of the subtypes
found by Fujino. Note that cells 4-1 and 5-6, while reacting essentially negatively with horse
serum, were still capable of removing some of the antibodies or other erythrocytes reacting withthis serum.

observed were due to the coating of erythrocytes with Ju2 SSS. One experimentof this kind was performed by placing drops of washed erythrocyte suspensions fromtwo Ju2-negative whales (10-2 and 11-7) in drops of undiluted serum from aJu2positive whale ( 11-11). These mixtures were allowed to stand for twenty minutes,
at the end of time which ± traces of agglutination attributed to agglutinins otherthan those involved with the Ju system were observed. No agglutination occurredin saline controls of the cells or in the mixture of 11-11 cells with their homologous
serum.

74

J.E. CUSHING, K. FUJINO AND N. CALAPRICE

Drops of horse ·serum ( 1 : 5) were then added to the three mixtures of cellswith 11-11 serum. At the end of a second period of twenty minutes cells of 10-2
and 11-7 showed strong (tit/) agglutination while those of 11-11 were negative. In
contrast horse serum placed in the saline controls gave only a ± agglutination with
10-2 cells, none with 11-7 cells, and tit/ with 11-11 cells. These results canbe interpreted as due to the coating of the erythrocytes with Ju2 SSS in the case ofcells I 0-2 and I I- 7, and to the inhibition of horse serum in the case of 11-11. They
were confirmed in a second series of experiments that showed the titer of Ju2 SSS
inhibitor in 11-11 serum to be one in sixteen with respect to the reactions of horse
serum with 11-11 cells; and one in four with respect to ability to coat the cells of
11-7. (The serum of 11-7 contained strong iso-antibodies for 11-11 cells and for
all the Ju2-positive cells shown in table 5. It did not inhibit the reactions of horse
serum with these cells).
A second experiment similar in kind to the one just described was performedby placing a suspension of human type A cells in the serum of anotner Ju2-positive
whale 5-3. This serum was also known to inhibit the agglutination of its homologous cells by horse serum. No agglutination of the human cells occurred in thisserum after twenty minutes. The adition of horse serum (one in ten) to the
mixture again resulted in a strong agglutination after a twenty minutes wait. The
cells of Ju2-negative whale 0-2 also were affected by 5-3 serum in the same way.
Controls showed that the horse serum alone gave only traces of agglutination withthe human cells and none with the 5-3 whale cells.
A third experiment was made with the serum of Ju2-positive whale 0-1 also
known to contain horse serum inhibitor and not to contain isoantibodies. In
this case, however, it was not possible to induce the agglutination of human A
cells or those ofJu2-negative whale 0-2. The serum of whale 0-2 did not appear tocontain Ju2 SSS as shown by an inhibition experiment which took into accountthe presence of isoagglutinins in the 0-2 serum.
One set of experiments was made which showed that the serum of Ju2 positive whale II-II strongly inhibited the agglutination of its own cells by finback
anti-Ju2 serum (no. 34) as well as that of the cells of Ju2 positive whale 0-1.
DISCUSSION

While alternative explanations can not yet be ruled out, the observations reported
above support the concept that the changes in type of sperm whale erythrocytes
were the result of post-mortem coating withJu2 SSS. Changes of a similar nature
have been produced experimentally with the cattle J antigen, originally reported by
Stormont (1949) and more recently by Stone (1962). It is also of some interest
that Stone et al (I 954 p. 399) comment that occassionally a sample of cattle cells
classified as J negative later showed very weak reactions with high concentrations ofJ reagents, and that these individuals occurred among those whose serum contained J inhibitor. Experiments reported by Kodani (1962) show that the bloodgroup phenotypes of human epithelial cells can also be altered by exposure toJU BLOOD TYPING SYSTEM AND SOLUBLE SUBSTANCES

75

soluble heterologous blood group substances. The occurrence of Ju2-negative
whales whose serum contained isoantibodies for Ju2-positive cells, and which were
not capable of inhibiting the action of horse serum, show that the two major classesof whales found in California and the Antarctic also exist in the Peruvian population. However, the frequency of individuals whose erythrocytes reacted strongly
with horse serum (including the five that changed type) was much higher in the
Peruvian sample than in the others. Such whales totaled 25 among a sample of 33
whales that were systematically typed with horse serum and the two antifinback heteroimmune serums. (Whales 4-1 and 5-6 are counted as negative in this instanceas they did not agglutinate in horse serum).
The tentative explanation offered for the high frequency of Ju2-positive
erythrocytes in the Peruvian material is based on the fact that the blood was obtained
from whales in a relative advanced post-mortem condition due to a longer period,in higher temperatures of sea and air, between death and £lensing. In addition,
the samples, having been taken from supercial cuts rather than the heart, were
exposed to mixture with body fluids other than serum. These factors could wellprovide a reasonable opportunity for Ju2 SSS to coat the erythrocytes after thedeath of the animal. If this explanation is correct the Ju2 antigen, by comparison
with Antarctic data, would seem to occur on the erythrocytes of living individuals
with much less frequency than it occurs in the serum and or other tissues as a soluble
substance.
The above considerations obviously lead to an evaluation of the usefulness of
the Ju antigens in population reaserch. Two points are to be made. First, thephenomenon of type change was not observed by Fujino in his studies on sperm
whales. This supports the view that fresh blood samples taken from the heart
present an accurate picture of the relations between J u2 antigen in the serum andattached to erythrocytes. Second, no type changes have been observed in the very
extensive research that has been done on finback whales. Here not only are theconditions of sampling significant, but also the fact that this species, being a baleen
whale, is very far removed in an evolutionary sense from the sperm whale. Thisis significant as it is known that the human ABO substances vary with respect totheir solubility and occurrence upon erythrocytes among the many species where
they occur (see, for example, Mourant 1954 page 175 on the primates). Other
points could be made, but enough has been written upon which to base two conclusions from the results of this paper. First, the Ju2 finback-like antigens insperm whales appear to be soluble antigens with properties akin to the soluble antigens known in other species. Second, samples taken from the heart blood of tecently killed whales appear to present valid material for frequency determinations.
However, further blood typing studies involving the Ju system will have to consider the possible complexities introduced by soluble components of this system andmethods of sampling. That the frequency of the different phenotypes producedby systems involving soluble antigens can be determined is already shown in them
literature on the occurrence of cattle J in various breeds (Sprague 1958, Stormont
1959) and by the studies that have been made on the human Lewis system (Race76

J.E. CUSHING, K. FUJINO AND N. CALAPRICE

et al. 1958).
SUMMARY

This paper reports the discovery of a pair of antigens in sperm whales that resemble
the Jul and Ju2 antigens of finbacks and other species. Reagents for the detectionof these antigens include heteroimmune antifinback serums, the normal serums ofseveral domestic animals, and isoantibodies for Ju2 in sperm whale serum. The
Ju2 antigen exists as a series of at least four subtypes. The distribution of Ju2positive individuals was determined in Antarctic samples of sperm whale populations. Observations on sperm whales taken off Peru suggest that the Ju2 antigen
can exist as a soluble substance which can cause changes of type in erythrocyteson post-mortem.
ACKNOWLEDGEMENTS

The authors acknowledge support and assistance from the following organizations
and persons: The Ministry of Education, The Japanese Government; The UnitedStates Office of Naval Research, Biology Branch (Contract No. NONR 03503);
the officers and the crews of the factory ship, Kyokuyo-maru No. 2; The Cia
Balleneria Del Norte, Tierra Colorado, Peru; The Del Monte Whaling Co.,San Francisco; The Golden Gate Whaling Co., San Francisco; Mr. Dale Rice,
U.S. Bureau of Commercial Fisheries, Seattle; and Dr. H. Omura, Director, The
Whales Research Institute, Tokyo. Dr. Lucian Sprague, U. S. Bureau of Commercial
Fisheries, Honolulu, made valuable suggestions concerning the preparation ofthis manuscript.
LITERATURE CITED
CUSHING, J.E. (1962). Immunogenetic concepts in marine population research. American Naturalist,(889):CUSHING, J.E., K. FUJINO and K. TAKAHASHI. (1959). Glycerol-freezing technique as an aid in blood typingof whales. Sci. Rep. Whales Res. Inst. 14: 89-100.
FUJINO, K. (1954). On the serological constitution of the sperm- and baird beaked-whales (I). Bloodgroups of the sperm and baird-beaked whales. Sci. Rep. Whales Res. Inst. 9: 105-120.
FUJINO, K. (1960). Immunogenetic and marking approaches to identifying subpopulations of the North
Pacific Whales. Sci. Rep. Whales Res. Inst. 15: 85-142.
KODANI,, M. (1962). In vitro alteration of blood group phenotypes of human epithelial cells exposed toheterologous blood group substances. Proc. qf Society for Experimental Biology and Medicine, 109 : 252-258.
MouRANT, A. E. (1954). The distribution of the human blood groups. Charles C. Thomas Publ. Springfield,
Ill.
RACE, R.R. and R. SANGER. (1958). Blood groups in man. 3rd ed. Blackwell Scientific Publ. Oxford.
SNEATH, J. S. and P.H. A. SNEATH. (1959). Absorption of blood-group substances from serum on to red
cells in Blood Groups. British Medical Bull. 15: 154-157.
SPRAGUE, L. M. (1958). On the recognition and inheritance of the soluble blood group property "Oc"of cattle. Genetics, 43: 906-912.
STONE, W. H. (1962). The J substance of cattle. Annals of the New York Academy of Sciences, 97 (I) : 269-280.

JU BLOOD TYPING SYSTEM AND SOLUBLE SUBSTANCES

77

STONE, W. H. and M. R. IRWIN ( 1954). The J substance in cattle I Development and immunogenetic studies.
]our. Immunol., 73: 397-414.
STORMONT, C. (1949). The acquisition of the J substance by bovine erythrocytes. Proc. National Academyof Sciences, 35 : 232-237.
STORMONT, C. (1959). On the applications of blood groups in animal breeding. Proc. X Int. Congress ofGenetics
I, 206-224.
STORMONT, C. (1962). Current status of blood groups in cattle. Annals of the New Tork Academy of Sciences,97(1): 251-268.
STORMONT, C., Y. SuzuKI and W.J. MILLER (1960). Bacterially mediated false-positive reactions in lytic
blood-typing tests. Nature, 186 (4720): 247-248.
STORMONT, C. and Y. SuzuKI. (1960). On the "J" classification of rabbits and the production of anti-Jin" ]-negative" rabbits. Proc. of the Soc. for Exp. Biol. and Med., 105: 123-126.