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324 



BLOOD TYPES IN PACIFIC SALMON 



INIarTne Biological Laboratory! 

1960 
WOODS HOLE, MASS« 




SPECIAL SCIENTIFIC REPORT- FISHERIES No. 324 




UNITED STATES DEPARTMENT OF THE INTERIOR 
FISH AND WILDLIFE SERVICE 



SPECIAL NOTE 



The International North Pacific Fisheries Commission, 
established in 1953 by the Internationjil Convention for the 
High Seas Fisheries of the North Pacific Ocean, coordinates 
the research of the member nations: Japan, Canada, and the 
United States. The resulting investigations provide data to 
the Commission for use in carrying out its duties in connec- 
tion with fishery conservation problems in the North Pacific 
Ocean. Publication of this scientific report has been 
approved by the United States Section of the Commission. 



United States Department of the Interior, Fred A, Seaton, Secretary 
Fish and Wildlife Service, Arnie J. Suomela, Commissioner 



BLOOD TYPES IN PACIFIC SALMON 



by 



George J. Ridgway Eind George W. Klontz 

Fishery Research Biologists 

Bureau of Commercial Fisheries 



Contribution No. 12 to research conducted with 
the approval of the United States Section of the 
International North Pacific Fisheries Commission. 




United States Fish and Wildlife Service 
Special Scientific Report — Fisheries No. 324 



Washington, D. C. 
January 1960 



11 



TABLE OF CONTENTS 

Page 

Introduction 1 

Material and methods 3 

Results 3 

Discussion 6 

Acknowledgments 7 

Summary 7 

Literature cited 7 



111 



BLOOD TYPES IN PACIFIC SALMON 



by 

George J. Ridgway and George W. Klontz 

U. S. Fish and Wildlife Service 

Seattle, Washington 



ABSTRACT 



Intraspecific differences in erythrocyte antigens (blood types) were shown to occur in four species 
of Pacific salmon, the sockeye or red salmon ( Oncorhynchus nerka ). the chinook or king salmon (O. 
tshawvtscha) . the chum salmon (O. keta) , and the pink salmon (O. qorbuscha ) ■ Antisalmon-erythiocyte 
sera prepared in rabbits and chickens were used after absorption of species-specific antibodies. Some 
of these blood types were shown to differ in their frequency of occurrence between different geographic 
races. In addition, isoimmunizations were conducted on one race of sockeye salmon. Antisera of 
seven different specificities were prepared and at least eight different patterns of antigenic composition 
were displayed by the cells tested. 

These results indicate that considerable antigenic diversity exists in salmon. Reagents to detect 
valuable markers for the investigation of geographic races of salmon should be obtained through further 
research. 



INTRODUCTION 

One of the most importcint problems in 
fishery biology is recognition of "races", 
or reproductively isolated subpopulations , 
of fishes. Since there may be little or no 
interbreeding or recruitment between such 
populations, successful regulation or 
management of fishery resources must be con- 
ducted in such a way as to recognize their 
independence. 

Present methods for distinguishing 
between the races of a particular species 
of fish are based on tagging by various 
means, or by seeirching for morphometric or 
meristic differences between the races 
(Rounsefell and Everhart 1953). 

It has been shown with many animals 
(Mayr 1942, Dobzhansky 1951), including 
fishes (Gordon 1947), that reproductively 
isolated subpopulations of species differ 
in the frequencies of one or more varieible 
genes. Therefore, a very useful adjunct to 
the methods of racial identification, now 
used in fishery management and research, 



would be the recognition and definition of 
genetically controlled polymorphic charac- 
ters. One class of such characters which 
has been found in every animal adequately 
investigated is intraspecific antigenic 
variations in the red blood cells. These 
characters are more simply known as blood 
types. 

Lcumdsteiner (1900) was the first to 
demonstrate the occurrence of blood types. 
He found them in man with natural isoagglu- 
tinins. In the same year Erlich and Mor- 
ganroth (1900) demonstrated blood types in 
goats using immune isohemolysins induced 
by transfusions. Subsequently, many addi- 
tional instances of intraspecific antigenic 
differences have been demonstrated. The 
results are most notable in the case of man 
(Race and Sanger 1954), cattle (Stormont 
et al. 1951) and chickens (Briles et al. 
1950). 

In each of these species, blood types 
have been used to study reproductively 
isolated populations. Thus, blood type 
frequencies have been used to characterize 



races of men (Boyd 1950, Mouremt 1954), 
breeds of cattle (Owen et al. 1947) and 
inbred lines of chickens (Schultz and 
Briles 1953). 

The studies of Fujino (1958) demon- 
strating the occurrence of blood types in 
whales are of particular interest to marine 
biologists. 

Until recently there have been rela- 
tively few published studies concerning the 
possible existence of blood types in fishes. 
Some of these demonstrated marked interspe- 
cific differences in cellular antigens but 
provided little or no evidence for intra- 
specific heterogeneity of such antigens. 
Noguchi (1903a, b) found that the serum of 
several species of fishes would agglutinate 
the red blood cells of other species but 
no intraspecif ic differences were noted. 
He did produce isoagglutinins and isohemoly- 
sins in two species of turtle, Chrysemys 
picta and Chelopus guttatus , demonstrating 
that cold blooded vertebrates may possess 
intraspecif ic antigenic differences, Toth 
(1932) found no evidence for the existence 
of blood types in carp from 280 cross- 
matches. Jensen (1937) tested for natural 
isoagglutinins in the cod ( Gadus morrhua ) 
and also attempted to demonstrate individual 
differences with the serum of a single 
rabbit immunized with the red cells of an 
individual cod. He did not find any conclu- 
sive evidence for the existence of blood 
groups in cod. 

Suyehiro (1949) reviewed the previous 
Japanese literature in which there were 
no instances of the demonstration of blood 
types in fishes. He was evidently the 
first to demonstrate individual antigenic 
differences in fish blood cells since he 
found a few instances of natural isoagglu- 
tinins in the eel ( Anguilla japonic a ) and 
the gilthead ( Sparus swinhonus Gimther). 
In 249 crossmatches of blood cells and 
serum from cod ( Gadus macrocephalus ) he 
found no evidence for natural isoaggluti- 
nins. Suyehiro also tested blood samples 
from 336 fishes of 30 different species 
with human ABO blood typing sera and found 
that only 91 were agglutinated. In 21 of 
the 30 species, individuals varied in their 
reactivity to the human sera tested. 

Gushing and Sprsigue (1953) studied 
the agglutinative activity of human anti-A 
and anti-B sera and rabbit antisheep cell 



serum for the erythrocytes of a number of 
species of fish. Considerable £intigenic 
diversity was noted between species but no 
individual differences were found within 
the members of the species tested. 

Recent reports have indicated a major 
breakthrough in the efforts to discover 
blood types in fishes. Hildemann (1956) 
applied the method of isoimmunization, 
which has been so successful in detecting 
blood types in other animals, to goldfish 
( Carassius auratus ) , One of the isoimmune 
sera detected six different antigenic types 
of goldfish. Immune rabbit sera were also 
prepared by Hildemann, one of which de- 
tected a single antigenic difference after 
careful absorption. Gushing (1956) re- 
ported the existence of individual anti- 
genic differences in the oceanic skipjack 
( Katsuwonus pel amis Linnaeus) detectable 
with natural isoagglutinins, normal bovine 
serum, and the sera of rabbits immunized 
with the whole blood of oceanic skipjack, 
albacore (Germo alalunga Gemlin), or 
Pacific mackerel ( Pneumatophorus japonicus 
diego Ayres), Gushing and Durall (1957) 
discovered and analyzed a natural isoagglu- 
tinin system in the brown bullhead (Icta- 
lurus n, nebulosus Le Sueur). This system 
was found to be analogous to the human ABO 
system in that four antigenic types were 
found (i.e., some fish possessed antigen 1, 
some antigen 2, some both antigens, and 
some had neither), and when an antigen was 
lacking, its corresponding isoagglut inin 
was always present. Ridgway, Gushing, and 
Durall (1958) found quantitative differ- 
ences in the reactivity of the cells of 
individual sockeye salmon ( Oncorhynchus 
nerka Walbaum) with natural antibodies from 
pig sera and demonstrated that there were 
significant differences between geographi- 
cally separated populations in the frequency 
of the different types detected. 

Suzuki et al. (1958) have demonstrated 
the existence of blood groups in species of 
tunas amd have presented evidence which 
suggests that differences exist between the 
blood type frequencies of albacore from the 
Pacific and Indian Oceeins. 

The present report provides further 
evidence for the existence of blood types 
in Pacific salmons and additional evidence 
that some of these characters differ in 
their frequency of occurrrence in different 
races of the same species. 



MATERIALS AND METHODS 

Most blood samples were taken in the 
field by severing the caudal artery with 
a sharp knife and collecting the spurting 
blood in sterile bottles. Some of the 
samples, including those used in isoimmuni- 
zations, were taken by cardiac puncture. 
The clotted samples were maintained on ice 
or in a refrigerator until used. Samples 
excessively hemolysed or over ten days old 
were not used for testing. Blood cells 
from samples as old as three weeks were 
used for animal inoculation. Cells for 
testing were washed three times in 10 to 50 
volumes of modified Alsever's solution 
(Bukantz et al. 1946) and adjusted to a 2- 
percent concentration in this solution. 
The use of the Alsever's solution as a 
suspending medium was found to be essential 
since salmon red blood cells lysed in saline 
or phosphate-buffered saline solutions. 

Most antisera were prepared by giving 
intravenous or intraperitoneal injections 
of 0.5 cc. of a 50 percent suspension of 
washed cells. Such injections were given 
to rabbits and salmon three times a week 
for three weeks and to chickens every three 
days for three or four injections. Four 
to twelve days after the last injection the 
cuiimal was bled and serum collected. In 
most cases, in order to detect individual 
differences, several additional stimulations 
at intervals or two weeks to a month were 
required. Some rabbits also received sub- 
cutaneous inoculations of washed suspensions 
of particulate material from lysed red 
cells in Freund's adjuvcmt. The sera were 
preserved by freezing and stored at -30° 
C. Serum dilutions were made with phos- 
phate-buffered saline or 1 percent saline 
solutions. 

Absorptions were performed by mixing 
a 1/2, 1/5 or 1/10 dilution of the heat- 
inactivated antiserum with an equal volume 
of washed packed cells, incubating for one 
hour at room temperature or in the refri- 
gerator, centrifuging the cells down and 
decanting the absorbed serum. Usually more 
than one absorption was required to remove 
all of the antibody present which would 
react with the absorbing cells. 

Tests were performed by mixing 0.1 ml, 
of absorbed serum, diluted if necessary, 
with 0.1 ml. of 2 percent cell suspension 



Table 1, — Individual antigenic differences demonstrated 
by agglutinin absorption tests in the erythrocytes 
of sockeye salmon from Cultus Lake and Adams River, 

(Rabbit antisockeye salmon-erythrocyte serum R19 absorbed 
and cross-tested with the cells of individual salmon) 







Test 


cells 




Absorbing 




Cultus Lake 


Adams River 




cells 


1 


15 19 12 13 


15 11 « 13 


8 



Cultus Lake 1 
Cultus Lake 15 
Cultus Lake 19 
Adams River 15 
Adams River 11 
Adams River 9 
Adams River 13 
Saline control 



+ 


+ 





+ 





+ 


+ 


+ 


+ 


+ 











+ 














+ 












































+ 

















































































in Alsever's solution in 10 x 75 mm. tubes. 
Dilutions are expressed as the final dilu- 
tion of the original serum, taking into 
account the dilution by the red cell sus- 
pensions. Readings were made after suit- 
able incubation periods at room temperature 
and usually after overnight incubation in 
the cold. The settling pattern was judged 
either smooth (S) or rough (R) and the 
degree of agglutination scored as 0, +_, 1 
plus, 2 plus, 3 plus and 4 plus. In order 
for a test to be considered, the saline 
control had to be smooth and negative. 



RESULTS 

The major part of our effort has been 
directed toward finding blood types in red 
or sockeye salmon ( Oncorhynchus nerka 
Walbaum). Toward this end we have immunized 
26 rabbits and 20 chickens with the washed 
erythrocytes or stroma from members of this 
species. For each of the other species of 
Pacific salmon we have made 2-8 rabbit and 
chicken anti-erythrocyte sera. Individuals 
or pools from individuals from the same 
area were used. These antisera were ab- 
sorbed with the erythrocytes of individual 
salmon from other areas and the resulting 
sera tested for residual activity for the 
cells of a number of individuals. With most 
antisera produced, numerous absorptions did 
not reveal any evidence for antigenic het- 
erogeneity within this species. However, 
blood group differences were demonstrated 
with a few rabbit immune sera. One of these 
sera (R19) was analyzed by absorption with 
the cells of sockeye salmon collected in 
1955 from Cultus Lake and the Adams River, 



Table 2. — Serological differences between races of 
Fraser River sockeye salmon. 





Number 
tested 


Number reacting wi 
prepared by absorb 
R19 serum with 


th reagents 
ing rabbit 
cells of 


Number not 
reacting 


Area 
(test 
cells) 


Cultus 

Lake 

19 


Cultus 
Lake 

1 




Cultus 
Lake 

15 


with any of 
the three 
reagents 


Cultus 
Lake 

Adams 
River 




17 
17 


3 
11 


2 
17 




12 
16 


4 



Chi-square 




7.6* 


56.6* 




3.3 


4.5* 



* Significantly different at the 95 percent level. 

both tributary areas on the Fraser River 
in British Columbia (table 1). The three 
samples tested from the Adams River race 
removed all or nearly all of the antibodies 
present in the serum. The three samples 
from the Cultus Lake race each removed all 
of the antibodies specific for their anti- 
gens, but left cintibodies which would react 
with cintigens present on the cells of other 
individual tested. Thus, three reagents 
of different specificites were pro- 
duced. 



Chinook salmon from the Columbia 
River, resulted in the production of 
reagents of four different specifici- 
ties. The results indicate a charac- 
teristic is present in number one 
which is absent from the other three. 
Superimposed on this, there appears 
to be a kind of subtype difference 
between 2, 3, and 4 which is shared 
with 1. 

Evidence for blood groups in 
chum salmon (0. keta ) is presented in 
tables 4 and 5. The experiments out- 
lined in these tables were performed 
with samples from the Samish River in 
Washington State. Evidence for the 
presence of a considerable amount of 
antigenic heterogeneity in this species and 
this race is provided by the results pre- 
sented in table 4, since reagents of four 
different specificities were obtained by 
five different absorptions. 

Five patterns of reactivity were dis- 
played by the samples tested; numbers 1, 6, 
8, and 9 reacted with the sera absorbed by 
2, 4, and 5; numbers 2 and 7 reacted with 



These results as well as addi- 
tional tests made on sajnples col- 
lected from these areas in 1956 with 
the above absorbed sera are combined 
in table 2. Statistically signifi- 
Ccuit differences in the proportion 
of individuals reacting positively 
with these reagents are apparent for 
the two populations. 

Some of the antisalmon red 
blood cell sera, prepared in rabbits 
and chickens, were useful in demon- 
strating blood group differences in 
species of salmon other than the one 
used for immunization. This is 
somewhat analogous to the demonstra- 
tion of the Rh blood groups in humans 
by Landsteiner and Weiner (1941), 
through the use of antirhesus monkey 
red blood cell sera. 

The evidence for blood groups 
in Chinook or king salmon (0. tsha- 
wytscha ) is presented in table 3. 
Absorption of serum R13, an anti- 
sockeye salmon red blood cell serum, 
by the cells of four different 



Table 3. — Demonstration of individual antigenic differences 
in Chinook salmon erythrocytes using rabbit antisockeye 
salmon serum (Serum R13 absorbed and cross-tested with 
four individual samples of Chinook erythrocytes). 



Absorb] 


■ng 


ce 


11 


Tes 


t ce 


11 


Dilutions of 


absorbed 


sera 


sample 


numbe 


rs 


sample 


numbers 


1:10 


1:20 


1:40 


1 


:80 


1( 


a) 








1 
2 
3 

4 






























2 










1 
2 
3 
4 




++++ 

+ 
























3 










1 
2 
3 

4 




++++ 
+ 




+ + 
+ 


















4(b) 








1 




++++ 


++++ 


+++ 




+ 












2 




++++ 


+++ 




















3 




+++ + 


+++ 


++ 

















4 




+ 














(a) This absorbed serum tested against the cells of an 
additional 12 chinook salmon, none of which reacted. 
Positive reactions were obtained with the cells of 
12 different sockeye salmon. 

(b) This absorbed serum tested against the cells of 24 
additional chinook salmon. Definite reactions 
occurred with 16 of them. 



Table 4. 



the sera resulting from absorption 
by 1 and 5; number 4 reacted with 
the sera resulting from absorption 
with 1, 3, emd 5; number 3 reacted 
with sera from absorption with num- 
bers 1, 2, 4, and 5; and number 5 
reacted with the sera from absorp- 
tion with numbers 2 and 4. The re- 
sults obtained with individuals 2, 4, 
and 7 demonstrate some of the com- 
plexities involved in the antigenic 
differences between individuals of a 
single species. All gave the same 
reaction with the reagents prepared, 
except for the weak reaction of the 
cells of individual 4 with the re- 
agent prepared by absorption with 
the cells of individual 3. Consist- 
ent with this is the fact that reagents of 
identical specificity were prepared by 
absorption with cells of individuals 2 and 
4. In contrast, absorption by cells of 
individual 7 removed the antibodies reactive 
with the cells of all individuals tested, 
but the cells of individual 7 failed to 
react detectably in agglutination tests with 
reagents prepared by cibsorption with cells 
of individuals 2, 3, or 4. Such apparent 
deviations from the principles upon which 
absorption analyses are generally assumed 
to rest have been associated, in other spe- 
cies, with the multiple-allelic control of 
sets of related but nonidentical specifici- 
ties. The large amounts of cells required 
to absorb the antibodies present in this 
serum which would agglutinate the cells of 
all members of the species, precluded 
further investigation of this system. 



Absorbing 






cell sample 






numbers 


1 


2 


1 





+ 


2 


++ 





3 








4 


+++ 





5 


+++ 


+++1 


7 









-Demonstration of blood types in chum salmon 
using chicken antichinook salmon serum 

(Serum 202 absorbed and tested with 
cells of individual chum salmon) 



Test cell sample numbers 
3 4 5 6 7 

+ ++00 + 

++ ++++ ++ + 
+ 000 

++ +++ ++ 
++ +++ 



+++ +++ 



++ ++++ 



The results presented in table 5 are 
less complicated with apparently a single 
difference detected between chum salmon 
sample 18 and the others tested. Superim- 
posed on this however are rather marked 
differences in the strength of reactivity 
of the other cells tested which may indi- 
cate additional antigenic heterogeneity. 

Evidence for the existence of blood 
group differences in pink salmon (0. gor - 
buscha ) is presented in table 6. A single 
difference was noted which appeared to be 
correlated with the area of origin of the 
samples tested. However, positive state- 
ments about area differences must await 
analysis of a larger number of samples. 

In addition to rabbit 2ind chicken 
immunizations, extensive isoimmunizations 



Table 5. — Demonstration of blood types in chum salmon 
using chicken antisockeye salmon sera. 



Sera 



11 



12 



13 



Test cell sample number 
14 15 16 17 18 19 20 21 22 23 24 



Serum 210: 










Absorbing ce 


11 








No. 18 




+ 


++++ 


+++ 


No. 20 













No. 23 













No. 24 













Serum 220: 










Absorbing ce 


11 








No. 18 




++ 


++++ 


++ 


No, 20 













No. 23 













No. 24 














++ ++++ ++++ ++++ 









n- ++++ +++ +++ 









+++ + +++ +++ ++++ +++ 












++ 


+ 


++ 


++++ 


+++ 


++ 


































































Table 6. — Demonstration of blood groups in pink salmon 

(Antiserum chicken E213 absorbed with red blood 
cells from Cordova pink salmon #6) 



Sashin Creek 



Kodiak Cordova 



Sample number 
Reaction 



12 3 4 5 6 7 



13 4 5 9 



+ + + + + 



have been conducted on sockeye salmon 
(0. nerka ). The fish used in all of these 
experiments were from the Columbia River 
(where this species is known as the blue- 
back salmon). In 1955 and 1956, 20 to 30 
adult salmon migrating up the Columbia 
River were captured and isoimmunized over 
periods of one to two months without any 
detectable development of isoimmune anti- 
bodies. Since many of these fish died 
before an adequate period for the produc- 
tion of isoimmune antibodies had elapsed, 
no conclusions about the antigenic het- 
terogeneity of salmon could be drawn from 
these experiments. 

In order to have fish available for 
isoimmunization over a longer period, a 
group of sockeye yearlings were obtained 
from the Winthrop Hatchery of the U. S. 
Fish and Wildlife Service in 1956 and 
reared in salt water at the Deception Pass 
Marine Research Station of the State of 
Washington Department of Fisheries. Iso- 
immunizations involving 100 fish, 50 pairs 
being cross-immunized, were started in 
August of 1957. During the period of this 
experiment the water temperature ranged 
from 14° C, to 9° C. with a mean of appro- 
ximately 12* C. 

Of these 100 fish, 15 produced iso- 
immune antibodies after periods of 7 to 14 
weeks and 5 to 11 inoculations. Because 
of the small size of these salmon, due to 
their being reared in captivity, only small 
amounts of the isoimmune sera could be 
collected. In addition, the sera were of 
low titer, most reacting only to 1 in 4 
dilution. Sufficient amounts of serum were 
obtained from seven of these fish to com- 
pare the specificity of the reactions 
against a number of individuals. 

The results of these tests are pre- 
sented in table 7. Inspection of these 
data reveals that all seven of the sera 
possessed different patterns of specific- 



ity, and at least eight different 
patterns of antigenic composition 
were shown by the cells tested. Thus, 
it would appear that the extent of 
antigenic variability in salmon is of 
the same order of magnitude as that 
found in other animals which have 

^ been extensively studied. Further 

study and application of this hetero- 
geneity will depend on the availabil- 
ity of larger salmon, and experimental 

facilities for holding them. 



DISCUSSION 

The demonstration of blood group dif- 
ferences in four species of Pacific salmon, 
as outlined in this paper, along with the 
demonstration that, in some cases, these 
characters cEin serve as markers of racial 
identity, indicates the existence of valu- 
able tools for the solution of many of the 
population problems encountered in the 
mjuiagement and conservation of these impor- 
tant fishes. However, more research and 
developmental work must be done. One of 
the biggest problems is in the production 
of sufficient type-specific sera to test 
large numbers of individuals from popula- 
tions of interest. In all of the immune 
sera we have produced in rabbits and 
chickens, most of the antibodies reacted 
with antigens possessed by all members of 
the species or genus. The production of 
type-specific sera in even small amounts 
required large quantities of cells for 
absorption. 

It is interesting to note that sev- 
eral of the immune sera were capaible of 
demonstrating blood group differences in a 
species other than the one used for immuni- 
zation, but could not be used to demonstrate 
differences in the immunizing species. 
This would appear to indicate that related 
antigens are fixed in one species and 
segregating in another. These observations 
are somewhat analogous to antigenic rela- 
tionships which have been demonstrated 
between man and Rhesus monkeys (Landsteiner 
and Weiner 1941), and recently between 
cattle asid bison (Owen, Stormont and Irwin 
1958), and between tahrs ( Hemitragus jam - 
lahicus ) and a variety of other Artiodactyl 
species (Stormont and Suzuki 1958). Fur- 
ther extension of these observations in 
fishes may result in the production of 
type-specific sera in practical quantities. 



Our isoimmunization experiments 
with sockeye salmon indicate con- 
siderable antigenic diversity exists 
in this species. Similar Eind more 
extensive studies have been con- 
ducted with rainbow trout and will 
be presented in another paper. As 
with other animals, isoimmunization 
appears to be the most promising ap- 
proach to the problem of blood group 
differences in fishes. The species 
of Pacific salmon are particularly 
difficult animals on which to conduct 
isoimmunizations as they invariably 
die after becoming sexually mature 
at two to six or seven years of age. 
It is only shortly before their death 
(two weeks to three or four months) 
while they are on their spawning 
migration that they become readily 
available. They also grow quite 
slowly in captivity and only one species, 
the sockeye, can be readily held in fresh 
water throughout its lifetime. At least 
some of these difficulties can be overcome 
and we hope to continue our attempts to do 
so. 



ACKNOWLEDGMENTS 

The authors wish to express their 
appreciation to Drs, R. S. Weiser, J. E. 
Gushing and R. D. Owen for their valuable 
advice and encouragement and to Mary La 
Rocque and E. D. Ullman for excellent 
technical assistance. 



Table 7. — Comparison of the specificities of isoimmune 
sera produced in sockeye salmon. 



Serum 



Cells 



14-4 



14-6 



14-10 



15-1 



22-10 24-2 



13-1 








+ 


14-3 


+ 








14-5 





+ 





14-9 








+ 


15-1 


- 


+ 


+ 


15-7 











22-9 


- 


- 


- 


22-10 


+ 


+ 





24-1 


+ 


+ 






+ Agglutination, No agglutination. 



Not tested. 



13-2 



problems involved in its practical 
utilization are discussed. 



LITERATURE CITED 



BOYD, W. C. 

1950. Genetics and the races of man. 
Little, Brown and Company, Boston. 

BRILES, W. E. , W. H. McGIBBON, AND 
M. R. IRWIN 
1950. On multiple alleles affecting 
cellular antigens in chickens. 
Genetics, vol. 35, No. 6, pp. 633- 
652. 



SUMMARY 

1. Through the use of absorbed rabbit and 
chicken antisalmon-erythrocyte sera, 
blood types were demonstrated to occur 
in sockeye, chinook, chum, and pink 
salmon. 

2. Some of these types appear to differ in 
their frequency of occurrence between 
different geographic races. 

3. Isoimmunizations between individuals of 
race of sockeye salmon indicated the 
existence of at least eight different 
antigenic types or combination of types 
within this race. 

4. The usefulness of this kind of research 
to fishery management and some of the 



BUKANTZ, S. C. , C. R. REIN, AND 
J. F. KENT 
1946. Studies in complement fixation. 
II. Preservation of sheep's blood 
in citrate dextrose mixtures (modi- 
fied Alsever's solution) for use 
in the complement fixation reaction. 
Journal Laboratory Clinical Medi- 
cine, vol. 31, No. 4, pp. 394-399. 

CUSHING, J. E. 

1956. Observations on the serology of 
tuna. U. S. Fish and Wildlife 
Service, Special Scientific Report — 
Fisheries No. 183, 14 pp. 

CUSHING, J. E, AND G. L. DURALL 

1957. Isoagglutination in fish. Ameri- 
can Naturalist, vol. 91, No. 857, 
pp. 121-126. 



GUSHING, J. E. , AND L. SPRAGUE 

1953, Agglutination of the erythrocytes 
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