BLOOD GROUPS OF TURKEYS’
G. R. J. LAWZ, W. J. MILLER3, V. S. ASMUNDSON,
AND
C. STORMONT
Departments of Poultry Science and Veterinary Microbiology, University of California, Davis
Received September 21, 1964
LTHOUGH considerable information is available on blood groups of chick-
A ens, as reviewed by BRILES( 1962) and GILMOUR( 1962), relatively little information has as yet been published on blood groups of domestic turkeys
(Meleagris gallopavo). The present report, based mostly on the senior author’s
doctoral dissertation (1961 ), is concerned with that subject.
MATERIALS A N D METHODS
In any study of blood groups, there are two primary objectives: ( 1 ) the development of
antisera or typing fluids which will disclose intraspecific differences in red cell antigens, and
(2) the classification of those differences with respect to genetic systems of blood groups. I n the
present study, the first of these objectives differed little from the approach used in the studies of
blood groups in cattle and sheep in that both isoimmune and heteroimmune antisera were used
as sources of typing fluids. (For references to the literature on blood groups in other animals,
attention is called to the 1962 Conference on Blood Groups in Infrahuman Specie.)
Isoimmunizations: Preliminary isoimmunization of 47 turkeys resulted in the production of
seven antisera from which reagents for five blood factors named A, B. C, D and E were prepared.
Those reagents were then used to type a large group of turkeys from which donor-recipient
combinations were chosen for further isoimmunizations. The birds were paired in such a way
that some would he expected to produce anti-A, others anti-B, and so on. Some were paired in
such a manner that none of the known antibodies would be expected as, for example, the use of
a donor of type DE in immunizing a recipient of type ADE.
Because of their greater volume of blood and uniformly low level of serum lipids, male turkeys
were used almost exclusively as recipients. Although these birds were predominantly Broad
Breasted Bronze turkeys, some were Beltsville Small Whites and a few were F , hybrids of
Meleagris gallopavo and M . ocellata. Further descriptions of these strains and hybr:ds may be
found in papers by CARSON,
LORENZ
and ASMUNDSON
(1955). JOHNSON
and ASMUNDSON
(1957)
and LORENZ,
ASMUNDSON
and WILSON(1956).
Injections were made with 25 percent suspensions of washed r-d cells. In the first series of
isoimmunizations, the intervals between injections were 3 to 4 days. In the subsequent series, the
interval was lengthened to 7 days. The first injxtion (5 ml of cell suspension) was usually
intravenous. The second injection (5 ml) was intraperitoneal. The third and subsequent injections, all of 3 ml quantities, were intravenous.
Injections were usually continued only until the titer of isoagglutinins was sufficiently high
(at least 1:8) to warrant collection of antisera. Some birds from which antisera were collected
between one and two weeks were continued on the immunization schedule. However, if there
was no response after 8 weeks, the injections were discontinued.
All blocd was collected either in citrate solution (2 percent trisodium citrate plus 0.5 percent
NaC1) used in a ratio 1:4 or ALSEVER’S
solution used in a ratio of 1:l. T o remove fibrin, all
Supported in part by a grant (G6109) from the National Science Foundation.
’Present address: Basic Research Laboratory, IIy-Line Poultry Falms, Johnston, Iowa
Present address: Department of Genetics, Iowa State University, Ames.
Genetics 5 1 : L53-2til Fehruary 1965
254
G . R. J. LAW
isoimmune antisera were heated approximately one-half hour at 56"C, stored overnight at 4"C,
and then centrifuged to remove the precipitate (MILLER
1958). All blood samples and antisera
were collected from the right jugular vein in the manner described by LAW(1960).
Heteroimmunizations; All heteroimmune antisera were produced in adult, commercial rabbits.
Each rabbit received seven intravenous injections of 0.5 ml of a 20 percent suspension of washed
red cells from a selected dcnor over a period of 24 days. Antise:a were collected 10 days after
the last injection.
Blood-typing tests: All tests were saline agglutination tests performed at room temperature
(22 to 28°C) in 10 x 75 mm tubes using two drops (0.10 ml) of antiserum or typing fluid and
one drop of a 2 to 3 percent suspens:on of washed red cells. After the setup of the tests the tubes
were shaken and the first r2ading was made between 30 and 46 minutcs. The tubes were again
shaken and a seccnd reading was made usually 2 to 3 hours later. Although the microscope was
frequently used to check weak reactions, all recordings wer? on the bssis of degrees of macroscopic
agglutinition. Saline solution in a stringth of 0,009 was used in the washing and preparation of
cell suspensions and in diluting antisera and typing fluids.
Absorptions: For the absorptions isoimmune ant'sera were used undiluted or, at the most, 1:2.
Heteroiminune antisera were usually diluted 1:8 but som, were used in a 1:4 dilution. Serial
absorptions were performed using a ratio of one part washed, packed red cells to approximately
1.5 parts of antiserum per absorption. The absorptions were rcp-atzd (usually two or three times)
until all agglutinins for the absorbing bloods were completely exhausted.
RESULTS
Production of blood typing reagents: Excluding 47 birds used in the preliminary work, 92 turkeys were isoimmunized and 42 responded by producing detectable isoagglutinins, but only 26 birds produced antisera with titers of l :32
or higher. Generally, the response was very weak, the highest being 1:512 and
only two of 42 antisera reached that level. There was little or no difference in
response between those birds which received injections at 3 to 4 day intervals
and those which were injected at weekly intervals. There was, of course, considerable variation among birds in the time of appearance of isoagglutinins and
the time at which maximum titer was reached. The latter, however, averaged
approximately five weeks.
Attempts to reproduce certain of the isoantibodies by administering booster
injections some six months after the initial series were disappointing. Only two
of 15 birds which received booster injections responded with a titer of antibodies
equal to or greater than that of the initial series. Some of the birds were completely refractory to the boosters even after five weekly injections.
Relatively few of the isoimmune antisera proved to be useful as actual sources
of blood-typing fluids and only about one in five contained antibodies of expected
specificity with respect to blood factors A, B, C, D and E for which all donors
and recipients were typed. Although anti-D could have been expected in six out
of the 92 isoimmunizations and anti-E in seven, none of the recipients produced
either anti-D or anti-E. On the other hand, anti-A, anti-B and anti-C were rather
frequently produced.
In addtion to antibodies for blood factors A, B and C, antibodies for new blood
factors named F, G, H, I, J and K were obtained from certain of the 42 isoimmune
antisera. Antibodies of specificities either identical with or closely similar to antiA, anti-B and so on were obtained from rabbit antisera. We shall give further
BLOOD GROUPS O F TURKEYS
25 5
consideration to the various specificities after brief mention of the results of the
he teroimmunizations.
The results of the heteroimmunizations, which were limited to a total of 23
rabbits, were in marked contrast with all 139 isoimmunizations. Each of 20 of
the 23 rabbit antisera, with titers ranging from 1:512 to 1:8192, yielded on
appropriate absorptions one or more reagents useful in blood typing turkeys; and
most of those reagents could be used in considerably greater dilution than those
obtained from isoimmune antisera. Generally speaking, the agglutination reactions obtained with the rabbit reagents were more sharply defined than those
obtained with reagents from isoimmune antisera. Furthermore, in contrast with
certain of the isoimmune antisera, there was no marked diminution of titer on
storage at -20°C.
Among the antibodies (A, B, C, D, E, F, H and J ) which could have been expected in the heteroimmune antisera (i.e., from what was known of the blood
types of the seven donors in tests with isoimmune reagents). all except anti-E,
anti-F and anti-J were obtained. In addition, one reagent of a specificity distinctly
different from any of the isoimmune reagents was obtained from two of the 20
antisera. That reagent was named anti-L.
The A antibodies isolated from each of six rabbit antisera in which they were
expected were all of a specificity related to but not identical with the A reagents
obtained from isoimmune antisera. Although the rabbit A reagents reacted with
the majority of bloods designated as A-positive with isoimmune reagents, there
were certain A-positive bloods which they failed to agglutinate. In keeping with
subtyping conventions, the rabbit A reagents were designated A, and the isoimmune A reagents A,. A-positive birds whose red cells reacted with both anti-A,
and anti-A, were said to be of subtype A, whereas those whose red cells reacted
only with anti-A, were said to be of subtype AL.Ultimately it was shown by
absorptions that some of the isoimmune A reagents contained anti-A, subfractions.
Similarly, the C reagents obtained from rabbit antisera were designated anti-C,
whereas those from isoimmune sera were designated anti-C,.
Three B subtyping reagents were obtained from rabbit antisera. Two of these,
designated anti-B, and anti-B,, were not encountered in isoimmune sera but the
one designated anti-B, was of the same specificity as isoimmune B.
Also we note that two F subtyping reagents, anti-F, and anti-F,, were distinguished in studies of isoimmune F reagents but, as already indicated, there
was no evidence of anti-F in the three rabbit antisera in which F antibodies were
expected.
In all, 18 different reagents (including subtyping specificities) were developed.
Table 1 is a summary of the reactions of those reagents, excepting anti-E, with
the red cells of 17 turkeys selected to illustrate variety of types and subtyping
patterns. (The sources of the reagents are indicated in that table by means of
symbols interpreted in the footnotes.)
In Table 2 are shown the frequencies of each of the 18 blood factors in ten
lines of turkeys in the 1961 breeding flock. As may be noted. there were marked
256
G . R. J. LAW
TABLE 1
Reactions of the red cells of 17 turkeys with 17 blood typing reagents* (Bloods selected to
illustrate variety of types as well as subtyping relationships)
Reagents and sourcest
A, A,
Turkeys
366
396
415
856
857
879
1382
1393
1398
1451
2028
2166
2344
2496
2742
2796
2861
3
1
B,
B,
~
3
3
D E’,
C,
B, C,
2
1
F2 G H Hm I
2
1
1
1
3
2
J K
1
1
L
1
?
++
+
+ + +
+
+
+
+
+ + + ++ ++ + + + + + ++ +
++ ++ ++ ++ ++ +
+ + ++ + + ++
++
++
++ ++
+
+
++ +
++ ++ + + + ++ +++ +++ ++
+ + + + ++
++ + + + ++ ++ ++ + + ++
+ ++ + + + ++ + ++ ++ + + ++ + + +
Antieenic formulas
A,DHJL
A, B, H J L
A,B,DJ
B, C, F, H I J K
B, D H I J K L
B, C, D H L
A, D F, L
A, C, D F, 1
A, B, C, D F, I J
A, C, D I J
F,GJ
DHIL
D F,H I K L
GJ
A, B, C2 F, 1 J
A,DIKL
C,F,HG
The plus symbol ( f ) signifies a positive reaction with no attempt being made to indicate degrees of reaction whirl1
ranged from weak to complete clumping of the red cells. The dot symbol (.) signilies no agglutination.
t 1: Reagents obtained only from isoimmune antisera. 2: Reagents obtained only from heteroimmune (rabbitj antisera,
3: Reagents obtained both from isoimmune and heteroimmune antisera.
TABLE 2
Frequency of blood factors in ten lines of turkeys
Lines (number of birds typed in each line is shown in parentheses)
0.344
0.0
0.0
0.0
0.0
0.0
0.0
1.000
0.062
0.142
0.0
0.656
0.798
0.188
0.109
1.000
0.405
0.072
0.476
0.095
0.0
0.357
0.024
0.739
0.166
0.04a
0.0
0.523
0.095
0.667
0.428
0.905
0.581
0.116
0.303
0.116
0.0
0.395
0.046
0.768
0.140
0.023
0.0
0.325
0.791
0.487
0.163
0.628
0.296
0.0
0.318
0.091
0.159
0.341
0.0
0.522
0.205
0.0
0.0
0.636
0.660
0.364
0.273
0.818
0.591
0.0.
0.0
0.0
0.0
0.0
0.0
1.000
0.0
0.0
0.0
0.470
0.286
0.224
0.082
1.000
0.318
0.455
0.455
0.136
0.0
0.0
0.0
0.931
0.818
0.0
0.0
0.613
1.000
0.660
0.046
0.818
1.000
0.0
0.375
0.0
0.0
0.875
0.0
1 .ooo
0.0
0.625
0.0
0.125
0.750
0.375
0.0
0.125
1.000
0.0
0.0
0.0
0.0
0.0
0.0
1.000
0.0
0.0
0.0
0.0
0.0
0.040
0.0
0.960
0.750
0.0
0.0
0.0
0.0
0.333
0.083
1.000
0.250
0.083
0.0
0.292
0.791
0.625
0.042
0.542
0.256
0.077
0.051
0.051
0.0
0.128
0.102
0.486
0.51 1
0.026
0.385
0.308
0.692
0.540
0.051
0.205
BLOOD GROUPS O F T U R K E Y S
25 7
interline differences for most of the blood factors. Data were also available on
the frequencies of ten of the 18 blood factors in these same lines for the year
1960. There were no significant intraline differences for the two years.
used in the isolation
For further details on the
of the individual reagents
1960) using
Family studies: I n the
anti-A (or A,) and anti-H from isoimmune antisera, blood factors A and H
appeared as mutually exclusive alternatives; that is, individual turkeys had
either A or H, but never both or the absence of both. Family data indicated that
H-positive birds were homozygous recessive. Furthermore, it was stated that the
red cells of known heterozygotes did not react with anti-H but always reacted
with anti-A. However, the conclusion that anti-H did not react with the red cells
of heterozygotes proved to be incorrect. As subsequent studies showed, the red
cells of heterozygotes were capable of binding anti-H but simply failed to manifest the second stage reaction-in this case agglutination. Accordingly, the original H reagent and others like it were behaving as extreme “dosage” reagents;
that is, such anti-H reagents were capable of bringing about agglutination of red
cells possessing two doses of blood factor H but were incapable of bringing about
agglutination of those red cells with only a single dose. Subsequently, an H
reagent, here designated as Hm, was isolated from rabbit antiserum which was
capable of causing agglutination of the red cells of all birds with blood factor H.
The A system: Further studies showed that other blood factors in addition to
A and H belong to the A system. One of the first examples became apparent in
a family in which the male was heterozygous (aA,aAs)
f o r blood factors A, and A,
) blood factor H. As expected, the red
and the female was homozygous ( a H a Hfor
cells of all 12 offspring were agglutinated by the Hm reagent. Blood factors A,
and A, segregated in such a manner that there were six offspring with A, and
six with A,, but all six A, progeny possessed blood factor D which could have
been inherited only from the male parent whereas the six A, offspring were
D-negative. Further examples of the inheritance of blood factor D in association
with A, followed. (Also, it was noted that blood factor A, never occurred in the
absence of D.) Similarly, it was shown in certain matings that D segregated in
association with H. Thus, the two phenogroups AID and DH, presumed to be
controlled, respectively, by the alleles
became identified.
and aDH,
In the eight offspring from a mating of a male of phenotype A,B,DH (known
to be heterozygous for H) with a female of phenotype H, there were four offspring of phenotype A,B,H and four of phenotype DH. This suggested that B,
might be segregating with A, in a phenogroup A,B, controlled by an allele aAsBs.
Additional family data, in which the parents and offspring had not been classified
for subtypes of A and B and were not tested with the Hm reagent, showed that
blood factors A and B were segregating as a unit. Here, however, it was not
possible to indicate which subtypes of A and B were involved.
From a mating of a male of phenotype B,H with a female of phenotype
A,R,DH it was possible to classify the 14 offspring in four types: A,B, ( 3 ) , A2B,H
-
258
G.
R. J. LAW
TABLE 3
Inheritance of blood factors A and H
Number of progeny observed
Probable genotype
Mating+
Probable genotypes No. of matings
Number
1
2
3
4
5
6
7
8
9
aAaA x aAaA
aAaH X aAaA
anaA x nAaH
Total, No. 2 f No. 3:
aAaH x aAaH
aAaH X aH(aH)*
aH(aH)* x &aH
Total, No. 5 No. 6:
aH(aH)*X aAaA
aAaA X aH (aH) *
Total, No. 7 f No. 8:
aHaH x aHaH
+
a"&
6
2
3
5
8
8
2
5
-
6
25
10
10
20
13
0
0
0
0
0
0
0
a*aH
0
5
17
22
26
31
37
68
22
45
67
0
af[aJ[
Total
0
0
0
0
16
25
15
27
42
55
76
82
158
22
45
67
75
45
45
90
0
0
0
75
* Allele in parenthesis could have been a8 in certain matings since there were several offspring in each of the matings
5, 7 and 8 which were classified as genotype u4u'1 even though their red cells were nonreactive with H antibodies (see
text).
t Male parent listed first.
TABLE 4
segregation of alleles at the A locus of one male of phenotype A,B,D in mvrtings to three females
Dam
Mating
number
Phenotype
13
14
H
B2H
15
Totals
AID
Gametes from sire*
Gametes
aH
aB2
aH
Total
aAP
a*P
aAZB,
Total
6
6
2
3
5
12
4
8
12
8
32
2
5
7
5
3t
18
14
Number of progeny with phenotypes corresponding to union of gametes from sire and dam.
t Phenotype of these birds: 4B,D.
TABLE 5
Segregation of A alleles in progeny of mating 9 (a%aH x a*,%aDH)
*
Gametes from sire
Gametes
from dam
aDH
Totals
* Phenotypes
aBz
aH
Totals
3
3
3
6
6
8
14
5
8
of sire and dam were B,H and A,B,DH, respectively.
259
BLOOD GROUPS O F TURKEYS
( 3 ) , B,DH ( 5 ) and DH ( 3 ) , thereby suggesting that the male was of genotype
aBzarrand the female was of genotype aAjB*aDH.
Pertinent data relative to the aforementioned segregations are summarized
in Tables 3,4 and 5 .
In all, five A-system alleles aBe, aH,a"P, a 4 s B 1 and aD"were indicated in the
family studies. It is implied that other alleles involving the subtypes B, and B,
would have been identified had the reagents for those factors been available at
the time of the family studies. From this it also follows that certain birds (genotype 8,aBz)would be expected to be negative in tests for A and H. Indeed, such
birds were observed in two of the line, 630 and 690, in Table 2. Furthermore,
recent data (R W. C. STEVENS
1964, personal communication) have indicated
the existence of birds whose red celIs failed to react to all present A-system
reagents.
The C system: Family data (LAW1961) provided evidence that blood factor C
was segregating independently of those in the A system. It is proposed that blood
factor C (subtypes C, and C2) is a member of a second system of turkey blood
groups and that at least three alleles are involved in the control of that system.
Data on the inheritance of blood factor C in various matings are summarized in
Table 6.
The F system: Similarly, it was possible to show that blood factor F (subtypes
F, and FA)segregated independently of blood factors in the A system and also
independently of blood factor C (R. W. C. STEVENS).
There were, however, some
peculiar irregularities in the segregatibn of the subtypes of F, and F,. As an
example, among 14 progeny from a mating of an F-negative male (genotype 8 )
X an F,-positive female (presumed genotype f " l f ) , ten were classified as F1-positive and four as F-negative, as shown in Table 7. The difficulty here, and in other
matings, was that the parents were no longer available for retesting at the time
the offspring were typed. Thus, the possibility of misclassification of subtypes
could not be excluded. But there were other peculiarities concerned with the
distribution of the two subtypes. In four families all derived from matings of Fnegative males (ffl X F,-positive females
males of subtype F, exceeded
F,-negative males 16:7, whereas F,-negative females exceeded F,-positive females
18:8. However, the ratio 24:25 of F,-positive to F,-negative birds was in good
(fFlf),
TABLE 6
Segregation of cc alleles in progeny of mates possessing blood factor C
Number of progeny observed
Mating'
Genotypes
Probable
genotype
No of
matings
ccc x cc
cc x ccc
ccc x ccc
CVCC x ccc
3
* Male parent listed first,
4
3
1
~ C c Q a n ccc
d
16
24
21
6
cc
Totals
10
23
9
0
26
47
30
6
260
G. R. J. LAW
TABLE 7
Segregation of f F alleles in progeny of mutes possessing blood factors F , or F ,
Number of progeny observed
Mating+
Probable
genotype
ff x
4
7
f"1f
x ff
ff x f"J
x
fFd
f"f"2
No. of
matings
/"If
1
3
Phenotype
Fl
F*
24
0
37
0
I*
I*
lo*
24
No-F
Totals
25
42
49
80
4
0
25
14
* See text for a discussion of discrepancies.
+ Male parent listed first.
agreement with expectation (Table 7). (The chi-square test showed that the
deviation from the expected distribution of sex and phenotype F, was highly
significant.) However, no peculiarities of this order were noted in the distributions of blood factor F,.
Blood factors G, I , J , K and L: For one reason or another, the data were not
adequate to test the relationship, if any, of blood factors G, I, J, K and L with
those in the A, C and F systems, and with one another. In the case of G, as already
mentioned, the limitation was largely that of an inadequate number of G-positive
birds. In the case of I, it was the lack of I-positive x I-negative matings. The
majority of the matings involved parents which were both I-positive. In the case
of blood factors J and K it was largely the lack of sufficiently reliable testing
fluids. There was only one source of anti-J and the reagent was too weak, especially after prolonged storage at -20°C, to produce reliable results. Although
there were several sources of anti-K, all K reagents lost their strength rather
rapidly on storage at -20°C. Blood factor L was isolated too late to be used in
most of the family studies.
DISCUSSION
Most of the effort in the present study was devoted to the development of
sources of turkey blood-typing reagents from isoimmune antisera and much of
that effort, as we have seen, was to no avail. The majority of the birds were
simply refractory to the formation of detectable isoagglutinins. Other investigators [WOLFE
and DILKS( 1949) and WOLFE,
MILLER
and ASMUNDSON
(1961 ) ]
have noted difficulties in attempting to induce turkeys to form precipitins.
Response to isoimmunization is not 100 percent in other species and it is often
necessary to isoimmunize a considerable number of animals before the desired
antibodies to many of the blood factors are obtained in usable titer. Nevertheless,
in our experience the turkey seems to be particularly refractory to isoimmunization in contrast with experience in this laboratory in the isoimmunization of
cattle, sheep, chickens and horses.
One of the goals in these studies of blood groups in turkeys is to use the blood
group genes and loci as markers in studies of various quantitative characters
BLOOD G R O U P S O F T U R K E Y S
26 1
(production traits) to determine whether the blood group loci exercise any effects
on the latter traits as has been reported f o r certain blood group loci in chickens
1962):
(see review by GILMORE
S U MMARY
Data are presented on the production of 18 blood-typing reagents which
define individual differences in the antigenic structure of turkey red cells. Corresponding to the reactions obtained with those reagents, the antigenic factors A
(subtypes A, aqd A,), B (B,, B, and B?), C (C, and C,), D, E, F (F, and F2),
G, H. I, J, K and L were named.
Blood factors A, B, D and H were shown to be members of the same system,
a system named A, in which five heritable combinations (phenogroups) , namely
A,D, DH, A,B,, B2 and H, were identified. Each of the phenogroups is believed
to be controlled by one of the members of an allelic series of genes. Blood-typing
reagents of specificity H, as prepared from isoimmune antisera, were shown to
be extreme dosage reagents incapable of bringing about agglutination of the
bloods of H-positive individuals which had inherited an allele for H from only
one of their parents. The reagent named Hm, prepared from a rabbit antiserum,
was capable of bringing about agglutination of all H-positive bloods.
Blood factor C (subtypes C, and C,) is a member of a second genetic system
involving at least three alleles. Similarly, blood factor F (subtypes F, and F,) is
a member of a third system involving at least three alleles. Unexplained irregularities in the distribution and segregation of subtypes of F were encountered.
The data were not adequate to place blood factors G, I, J, K and L with respect
to genetic systems.
LITERATURE CITED
BRILES.W. E., 1962 Additional blood group systems in chickens. Ann. N.Y. Acad. Sci. 97:
173-183.
J. D.. F. W. LORENZ.
and V. S. ASMUNDSON,
1955 Semen production in the turkey male.
CARSON,
11. Age at sexual maturity. Poultry Sci. 34: 344-347.
Conference on Blood Groups in Infrahuman Species, 1962 Ann. N.Y. Acad. Sci. 97: 1-138.
GILMOUR.D. G., 1962 Current status of blood groups in chickens. Ann. N.Y. Acad. Sci. 97:
166-1 72.
A. S., and V. S. ASMUNDSON,
1957 Genetic and environmental factors affecting size of
JOHNSON,
body and body parts of turkeys. I. The heritability and interrelationship of body weight and
live body measurements. Poultry Sci. 36: 296-301.
LAW,G. R. J., 1960 Blood ssmples from jugular vein of turkeys. Poultry Sci. 39: 1450-1452.
1961 Blood groups of turkeys. Ph.D. dissertation, Univ. of California, Davis.
LAW,G. R. J.. and W. J. MILLER,1960 The A system of turkey bhod groups. (Abstr.) Genetics
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