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Blood groups of baboons. Population genetics of feral animals

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Blood Groups of Baboons. Population Genetics of
Feral Animals
W. W. SOCHA,' A. S. WIENER,' J . MOOR-JANKOWSKI AND C. J . JOLLY
'Primate Blood Group Reference Laboratory and World Health Organization Collaborating Centre for Primate Haematology, at the Laboratory for Experimental Medicine and Surgery in Primates (LEMSIP), of the New York University Medical Center,
New York,New York;3Department of Anthropology of the New York University, New
York,New York I0016
KEY WORDS Baboons Olive baboons Hamadryas baboons.
Human-type blood groups . Simian-type blood groups
ABSTRACT
Blood and saliva from 495 Ethiopan baboons were collected in
the field and tested for their human-type A-B-0 groups while 493 blood samples
were tested for their simian-type blood factors AP, Bp, Cp, GP, NP, ca and hu. Four
series of feral animals were tested: 194 olive baboons, a troop of 82 and another
of 90 hamadryas baboons, and a series of 129 baboons classified as olive/
hamadryas hybrids. In addition, 126 baboons from other sources were tested for
their human-type A-B-0 groups and 131 for their simian-type blood groups.
Human-type groups A, B and AB but not group 0 were found in combined
series of 621 animals. Gene frequency analysis also indicated the absence of
group 0. Population analysis of the data obtained for the 493 Ethiopian baboons
has shown that the simian-type blood groups AP and BP are independent of one
another. In contrast Bp and GP appear to be determined by corresponding allelic
genes; if confirmed by population data on additional series of animals, this would
define the first baboon blood group system found. There is a close association between the blood group specificities ca and hu, the exact nature of which still remains to be clarified. Blood group ca, originally believed to be species specific, is
found to be polymorphic in both olive baboons and hamadryas as well as in the
hybrids; hu, on the other hand, present in all hamadryas tested, is polymorphic
only in olive baboons.
Baboons, like other Old World monkeys and
apes, have homologues of the human-type AB - 0 blood groups (Moor-Jankowski e t al., '64;
Wiener and Moor-Jankowski, '69). In addition, baboons have their own, numerous, simian-type blood groups (Moor-Jankowskiet al.,
'67; Wiener et al., '701, analogous to the
multiplicity of blood groups in man and in
lower animals. Reagents for demonstrating
the simian-type blood groups of baboons have
mostly been produced by isoimmunization,
and also by cross-immunization (Moor-Jankowski et al., '73;Moor-Jankowski et al., '74).
Former attempts to classify the various simian-type agglutinogens of baboons into systems have been unsuccessful. This was due in
part to the unavailability of families for genetic studies and in part to the lack of large
AM. J. PHYS. ANTHROP., 47: 435-442
enough samples from normal feral populations to be used for methods of population genetic studies. Recently, we had the opportunity to test blood and saliva samples of large
series of baboons collected by F. L. Brett in
Ethiopia. Details of that field study are reported separately (Brett et al., '761, including
a troop by troop detailed analysis of the findings for the human-type A-B-0 group. The
purpose of this article is to analyze the blood
group findings further, applying the methods
of population genetics with special reference
to the simian-type blood groups.
MATERIAL AND METHODS
Baboons caught in the wild were subjected
' Supported by USPHS, NIH
Grant
GM 12074 and Contract HR
4-2184.
435
436
W. W. SOCHA, A. S. WIENER, J. MOOR-JANKOWSKI A N D C. J. JOLLY
TABLE I
Human-type A - B - 0 bloodgroups
Phenotypes
A
Population
EthiOpiQ
Hamadryas
troop A
troop K
combined
Olive
baboons
Hybrids
Subtotal
AB
B
Estimated gene frequencies
Tntal
NO.
No
'b
0
15
15
0.00
16.67
8.72
76 92.68
31 34.44
107 62.21
6
7.32
44 48.89
50 29.07
82
90
172
5
2.58
133 68.56
56 28.87
5
3.88
81
62.79
43
25
5.05
321
64.85
28
22.22
39
30.95
No.
%
No
%
Allelism
0
A
B
xz,t,
p
0.037
0.411
0.233
0.198
0.170
0.963
0.589
0.747
0.733
0.830
0.12
0.008
5.93
1.26
0.10
>0.70
>0.90
c0.02
>0.20
194
0.0
0.0
(a) 0.0
(b) 0.069
0.0
33.33
129
0.0
0.206
0.794
0.06
>0.80
149 30.10
495
>0.70
Other
sources
Total
53
360
59
46.82
208
to physical examination and taxonomic classification k f . Brett e t al., '76).Their blood specimens were collected both with and without
anticoagulant and their saliva samples were
immediately inactivated in a boiling water
bath, as described in a previous publication
(Moor-Jankowski et al., '64).The inactivated
saliva samples as well as the blood specimens
were stored in liquid nitrogen in the field and
then brought to New York City and stored
there in liquid nitrogen, Batches of blood were
thawed and reconstituted (Rowe e t al., '72)
immediately before testing in our laboratory.
The human-type A-B-0 groups were determined in 495 Ethiopian baboons by quantitative inhibition tests for A-B-H in the saliva,
and by testing the baboon sera for their content of anti-A and anti-B agglutinins after removal of non-specific heteroagglutinins by absorption with human group 0 red cells. Since
the introduction of this technique (Wiener
and Moor-Jankowski, '63) thousands of primate animals have been tested without difficulty and with readily reproducible results
confirmed by population genetics, family
studies (Moor-Jankowski and Wiener, '721,
transplantation of A-B-0 blood group compatible simian organs (Murphy e t al., '69) to
man and cross-circulation of human patients
with baboons (Fortner e t al., "71).
The simian-type baboon blood groups were
determined in all but two Ethiopian baboons
by testing the red cells with a battery of antisera, anti-Ap, anti-Bp, anti-CP, anti-GP, anti-
126
621
NP, anti-ca and anti-hu, described in detail in
earlier papers (Wiener et al., '70; Moor-Jankowski e t al., '73; Moor-Jankowski e t al., '74).
In general, these reagents produce their most
definitive reactions by the antiglobulin technique. The techniques of grouping simian
blood was reviewed by Socha e t al. ('72).
For the purpose of population analysis, the
Ethiopian baboons were divided into four
series on the basis of their physical characteristics and geographic location, namely, two
geographically widely separated groups of
hamadryas of 82 (Troop A) and 90 (Troop K)
animals respectively, a series of 194 olive
baboons (Papio anubisj and a series of 129 animals deemed to be olive/hamadryas hybrids,
giving a total of 495 animals. In addition,
there were 131 baboons (of which only 126
could be tested for their human-type A-B-0
blood groups) derived from a variety of other
sources; subspecies of those baboons had not
been determined.
RESULTS
In tests for human-type A-B-0 blood groups,
with only few exceptions, Landsteiner's rule
held, namely, there was a reciprocal relationship between the blood group substances present in saliva and the agglutinins present in
the serum, confirming the accuracy of the AB-0 group determination. As can be seen in
table 1, among the total of 621 animals tested
for the A-B-0 blood groups only groups A, B
and AB, but not group 0 were found. The fre-
437
BLOOD GROUPS OF BABOONS
quencies of the three blood groups in different
baboon troops were used for estimating gene
frequencies and for testing the hypothesis of
inheritance.
Table 2 summarizes results of tests performed on 493 blood samples of Ethiopian
baboons and 131 blood samples of baboons
from other sources, using baboon isoimmune
sera of seven different specificities. As can be
seen, t h e simian-type polymorphism was
found in all baboon troops for all specificities
tested except one, namely, hu, which was
present in all the 172 hamadryas baboons.
Because of the previously reported ambiguous observations on the relationship between
simian-type specificities AP and EP, the present findings on the relatively large sample of
animals are of special interest. Table 3, shows
distributions of the four types defined by antiAP and anti-Bp reagents, as well as the results
of Ap-Bp gene frequency analysis in different
series of feral baboons. The results obtained
are discussed in detail later in the article.
Table 4 shows the distribution of four blood
types defined by another pair of reagents,
anti-BP and anti-GP. Original observations on
simian-type specificities Bp and Gp suggested
the possibility of an antithetical relationship
between those two factors, and the present
series of baboons provided the opportunity to
test the possibility that Ep and GP might be
transmitted by triple allelic genes. The results of the gene frequency analysis are discussed later.
Finally, the table 5 presents the results of
tests with still another pair of reagents,
namely, anti-ca and anti-hu, which detect specificities supposed to be constrasting "species" characteristics of the red cells of two
groups of baboons: specificity ca of olive and
yellow baboons (cynocephalusanubis) and
specificity hu of hamadryas and chacmas
(hamadryas-ursinus). Discussion of the results of gene frequency analysis is deferred to
the latter part of the article.
Two-by-two contingency tests have been
carried out for associations between all pairs
of simian-type specificities other than Ap-Bp,
Gp-Ep and ca-hu. With only few exceptions the
tests indicated the absence of associations between any of the pairs of antigens analyzed.
In the few cases where the P values were less
than 0.5, the chi-square value is only just beyond the level of statistical significance.
Bearing in mind that when numerous 2 X 2
N O "
mml-2
m
$:
m
ooom
9999
m m
x
2
ooot-
m m
oooc
M
*
i
moYzz
N
O
N
-
m
wm
t -m
( occn
~
m m
2
m
Q,
3
N
m
w"2R
ri
*z
0
N
438
W. W. SOCHA, A. S. WIENER, J . MOOR-JANKOWSKI AND C. J. JOLLY
TABLE 3
Distribution o f A P - B p t y p e s
B l d types
op
Population
No.
%
Ethiopia
Hamadryas
troop A
troop K
combined
Olive
baboons
Hybrids
13
3
16
33
15.85
3.33
9.30
17.19
40
31.01
Subtotal
89
Other
sources
Total
49
138
np
AP
No.
%,
0
0
0
62
0.00
0.00
0.00
32.29
30
23.26
92
37.40
34
126
No.
9:
63 76.83
83 92.22
146 84.88
37 19.27
29
22.48
212
25.95
34
Gene
frequencies
APBP
Total
Ap
Bp
6
4
10
60
7.32
4.44
5.81
31.25
82
90
172
192
0.037
0.022
0.030
0.400
0.602
0.817
0.695
0.302
1.22
0.14
1.01
0.20
20.20
>0.50
>0.20
>0.50
30
23.26
129
0.269
0.263
0.82
>0.30
100
25.95
246
tests are carried out, even in the absence of
association one of twenty of the tests could be
expected to give results a t the borderline or
statistical significance, we do not consider
any of the seeming associations to be meaningful.
DISCUSSION
Significant differences in the distribution
of the human-type A - B - 0 blood groups have
been found for different series of baboons in
the present as well as in previous studies, beginning in 1964 (Moor-Jankowski et al., '64).
These differences in distribution appear to result from genetic drift caused by geographical
separation and isolation among the various
troops of animals. Thus, the A - B - 0 blood
group distribution in 192 olive baboons (P.
anubis) in the present study was different
from that observed in the previous series of
174 P. anubis but of different geographical
origin. Paradoxically, the distribution of the
A - B - 0blood groups in one troop of hamadryas
baboons in the present study does not differ
markedly from that of a group of olive
baboons from the previous study, but differs
markedly from that of the second troop of
hamadryas described here. These findings
demonstrate that for a single category of inherited markers, genetic drift can produce
wider differences among populations within a
species than those existing between species.
Not one group 0 animal was found among
the 621 baboons which represent the largest
number investigated by us in a single study;
Independence
rZili
X,
No
14
114
P
493
10.69
131
624
therefore, one might conclude that in this
series there were no carriers of the amorph
gene 0, so that only two allelesA and B had t o
be postulated. Therefore, these gene frequencies were calculated by direct count, and the
values obtained used to compute the expected
numbers of animals in each of three groups, A,
B and AB. The results of x 2 tests (cf. table 1)
indicate good fit for the 2-allele theory for the
population of olive baboons, for hybrids, as
well as for each of two troops of hamadryas,
when analyzed separately. Interestingly
enough, when the test for goodness of fit was
applied for the combined series of 172
hamadryas, the value of chi-square obtained
(xt = 5.93) seemed to reject the theory of inheritance based on the presence of only two
alleles. When, on the other hand, a different
method of estimating the gene frequencies
was used, this time based on assumption that
allele 0 is present but infrequent (Wiener and
Moor-Jankowski, '691, chi-square proved to be
only 1.26, showing a much better fit. Thus,
pooling of the two series with widely different
distributions of blood groups had resulted in a
misleading artifact. Thus, contrary to our previous assumption (Moor-Jankowski et al.,
'641, the presence of gene 0 in yellow, olive
and hamadryas baboons remains unproven.
On the other hand, however, two group 0 animals were actually found in a series of 188 P.
papio from Senegal (Wiener et al., '69);a third
group 0 baboon found in the same study was
of unknown origin and was not evaluated for
his morphological characteristics. It thus ap-
0 0.00
3
0 0.00
3
Oiher
sources
Total
0.00
1.56
0.00
0.00
Subtotal
0
0
0
3
X
No.
65.85
60.00
62.79
19.27
X
191
7
5.34
39 30.23
184
54
54
108
37
hu
No.
+-
0
ca
-+
10
%
7.75
157
107 81.68
50
TABLE 5
21 16.03
82
131
624
129
493
82
90
172
192
Total
++
34.14
40.00
37.21
58.33
Ib
273
17 12.98
256
80 62.01
28
36
64
112
No.
hulca
624
131
129
493
82
90
172
192
Total
0.098
0.156
0.126
0.084
0.577 0.263
0.571
0.748
0.641
0.291
0.722
1.0
1.0
1.0
0.527
0.450
0.566
0.640
0.600
0.5444
Ca
Gene
frequencies
Hu
@
6.12
0.125
0.831
0.30
3.12
0.05
0.02
6.5
4.7
-
P
-
Pill
XZili
12.1
8.93
27.8
XZIII
1.02
0.75
0.18
0.60
0.06
p
<0.001
<0.01
<0.001
P
Allelism
0.003
3.4
3.4
9.8
0.4
XtCH
-0.31
-0.88
r
0.95
0.07
0.07
<.01
0.55
p
Allelism
Chi-square for
Independence
Independence
0.160
0.331
0.096
0.233
0.625
@
Gene frequencies
Bp
Distribution of the specificities huand ca
24 18.32
76
61
11
8.53
X
27 20.93
52
No.
13 15.85
26 28.89
39 22.67
12 6.25
M
BPGP
3.66
2 2.22
5 2.91
20 10.42
3
No.
GP
0.00
0 0.00
0 0.00
40 20.83
0
No.
huica types
33 25.19
283
__
53 40.46
183
48 37.21
250
68.29
67.78
68.02
44.27
X
43 33.33
130
No.
Blwd types
56
61
117
85
%
BP
10 12.19
1 1.11
11 6.39
75 39.06
No.
OP
Ethiopia
Hamadryus
troop A
troop K
combined
Olive
baboon8
Hybrids
Population
Total
Other
sources
Subtotal
Ethiopia
Hamadryas
troop A
troop K
combined
Olive
baboons
Hybrids
Population
Distribution o f B P - G P t y p e s
TABLE 4
440
W. W. SOCHA, A. S. WIENER, J. MOOR-JANKOWSKI AND C. J. JOLLY
pears that while the distribution of the A-B-0
blood groups may vary greatly among as well
as within hamadryas, yellow and olive and
Guinea baboons, the presence of group 0,
though rare, has been demonstrated conclusively for Guinea baboons alone.
Anti-Ap and anti-Bp, the first two reagents
for typing red cells of baboons produced by isoimmunization (Moor-Jankowski e t al., '671,
have been studied most intensively. The fact
that both specificities are nonreactive or only
poorly reactive by the ficin method, as compared with the antiglobulin method, suggests
that the agglutinogens and antibodies involved are comparable to those of the human
M-N systems; this was one of the reasons why
we originally assumed that AP and BP belonged to one and the same blood group system. However, while no families were available for investigation, previous studies by the
method of population genetics on two series,
one of yellow baboons and the other of olive
baboons, indicated that AP and BP are independent of one another (Wiener et al., '70). On
the other hand, a different series of 184
Guinea baboons showed all four types, Op =
20, AP = 56, Bp = 12 and A p B p = 96. In that
case, it was evident that AP and BP exhibited a
rather strong positive association incompatible both with allelism and independence,
unless one invoked a fourth allele which produced an antigen having both specificities AP
and Bp, Thus, our previous results did not provide enough data to determine whether the
Ap-Bp baboon blood groups are transmitted by
allelic genes or are independent from one
another. The population genetics analysis performed in the present study leaves little doubt
that AP and BP are actually independent of
one another. As shown in table 3, the distribution of the Ap-Bp blood types in each of the
four series of baboons from Ethiopia demonstrates that AP and BP are independent from
each other, and, therefore, presumably belong
to different blood group systems. I t is instructive to observe the effect of pooling the four
series. When the statistical test for independence was applied to the combined series of
493 baboons, the very high chi-square value of
17.31 was obtained, making it appear as if
there were a strong negative association between the two agglutinogens AP and BP (r =
-0.19). Of course, this was an artificial association created by pooling several series of
baboons which produced a stratification
effect.
A relationship between agglutinogens Bp
and GP was suggested by the previous observation that reagents of specificities anti-BP and
anti-Gp could both be produced in baboons by
cross-immunization with red cells of geladas
fTheropithecus geladd (Moor-Jankowski e t
al., '71), even though anti-BP and anti-Gp sera
have been produced also by isoimmunization.
Though the possibility was not mentioned in
the original paper, we did notice that Bp and
GP appeared to be antithetically related, and
the present new series of baboons provided the
opportunity to test the possibility that BP and
GP might be transmitted by triple allelic
genes.
In combination, the two antisera, anti-BP
and anti-Gp,detect four blood groups (cf. table
4). The most striking results were obtained
with the series of 129 hybrids with the high
chi-square value of 6.12 for independence as
against only 0.003 for allelism. Thus, the distribution for the 129 hybrid animals conformed closely with the prediction under the
hypothesis of triple allelic genes for the BP
and GP blood groups, and argued against independence (P < 0.02). Also in the case of the
series of 192 olive baboons the findings favor
the hypothesis of triple allelic genes (x2(11=
0.4) rather than independence ( x * ( ~ =
, 3.12).
Paradoxically, however, for the series of 172
hamadryas baboons, the opposite results were
obtained, since chi-square for allelism was 9.8,
and for independence only 0.30. Still, it will be
recalled that the 172 hamadryas baboons consisted of two troops of animals, and when the
two troops were analyzed separately, the
paradox was again resolved since the chisquare for allelism was reduced from 9.8 to
only 3.4 for each troop. These data provide
still another example of the paradoxes which
arise when statistical analyses are applied to
series composed of a mixture of several different groups of animals.
In conclusion, therefore, the analysis of the
distribution of the Bp-Gp blood groups in three
series of feral Ethiopian baboons supports the
hypothesis that the resulting four blood types
are transmitted by triple allelic genes and not
by independent pairs of genes. This conclusion
is reinforced by the fact that these two factors, but not others, are shared by baboons
with geladas (Moor-Jankowski e t al., '74).
44 1
BLOOD GROUPS OF BABOONS
Therefore, if confirmed by further studies, the
resulting B p - G p system would represent the
first simian-type blood group system found in
baboons represented by more than a single
blood factor.
As mentioned before, the anti-ca and antihu reagents appear a t first to be “species specific.” If t h a t were so, the tests with these two
reagents would have shown only two contrasting types for ca and hu specificities, namely
+ - and - +. However, the tests carried out
on large numbers of baboons show that all
four combinations of these two specificities
can occur, namely
+ -, - and - (cf. table 5).
Of the four possible types, the rarest was
the one lacking both antigens hu and ca;
among the total of 624 baboons tested, only
three were found of that type (confirmed by
repeated tests). The existence of only three
major types,
-, - and
suggested
that the antigens hu and ca might be determined by contrasting allelic genes, but the
findings shown in table 5 do not support that
simple hypothesis. Thus, in two series, one of
olive and the other of hybrids, the frequencies
of the heterozygous hu-ca type were 58.3%and
62.0% respectively, whereas theoretically, the
frequency of that type could not have exceeded 50%if hu and ca were alleles in population
a t Hardy-Weinberg equilibrium. The distribution of the blood groups hu and ca among the
hybrid animals was intermediate between the
distribution for the hamadryas and olive
baboons, as to be expected. On the other hand,
the hybrid animals showed the highest frequencies of the heterozygous type hulca. All
the 172 hamadryas had the hu antigen, for
which the olive baboons and hybrids were
polymorphic. In contrast, baboons of all three
series were polymorphic for ca. In conclusion,
the findings indicate a negative correlation
between the hu and ca antigens, but the exact
nature and cause of this is not yet apparent.
Moreover, the results provide further evidence of the relationship between serological
specificities determining t h e species differences and those determining individual differences. For example, hu, shared by all animals of the two troops of hamadryas, proved
to be polymorphic in olive baboons and their
hybrids. On the other hand, specificity ca
which in our original study appeared to be
“species specific” for olive and yellow
+
+ +,
+
+
++
baboons, in the present study was shown t o be
polymorphic not only in hamadryas baboons
but also in olive baboons and their hybrids.
This result underlines the importance of testing large series of animals before concluding
whether a newly found serological specificity
is type specific or species specific.
We are aware that in the present study we
have applied to monkeys methodology originally developed for such studies in human
populations in which panmixia is assumed
(though it does not always occur) (Rosin, ’56;
Moor-Jankowski and Huser, ’56-57). The
breeding patterns of primate species vary.
Those of chimpanzee appear also to be different from those of man, yet the results obtained by applying human population genetics methodology to our blood group findings in
chimpanzees could later by fully confirmed
when families became available for testing
(Wiener et al., ’75). Similarly, we anticipate
that the conclusions presented in this paper
for baboons will be confirmed when baboon
families become available for blood grouping.
LITERATURE CITED
Brett, F. L., C. J. Jolly, J. Moor-Jankowski, W. W. Socha
and A. S.Wiener 1976 Human-type A-B blood groups in
wild Ethiopian baboons. Yearbook of Phys. Anthrop., in
press.
Fortner, J. G., E. J. Beattie, Jr., M. H. Shiu, J. S. Howland,
P. Sherlock, J. Moor-Jankowski A. S. Wiener 1971 The
treatment of hepatic coma in man by cross-circulation
with baboon. In: Medical Primatology 1970.Goldsmith
and Moor-Jankowski, eds. S. Karger, Baselmew York,
pp. 62-68.
Moor-Jankowski, J. K., and H. J. Huser 1956-57 Seroanthropological inveatigation in t h e Walser a nd
Romanch isolates in the Swiss Alps and their methodological aspects. Acta. Genet., 6: 527-531.
Moor-Jankowski, J., and A. S. Wiener 1972 Red cell antigens in primates. In: Pathology of Simian Primates.
R.N. T-W-Fiennes, ed. S. Karger, Baseli’New York, pp.
270-317.
Moor-Jankowski, A. S. Wiener and E. B. Gordon 1964
Blood groups of apes and monkeys. I. The A-B-0 blood
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Moor-Jankowski, J., A. S. Wiener, E. B. Gordon and J. H.
Davis 1967 Blood groups of baboons demonstrated with
isoimmune sera. Nature, 214: 181.
Moor-Jankowski, J., A. S. Wiener and W. W. Socha 1974
A new taxonomic tool. 11. Serological differences between
baboons and geladas demonstrated by cross-immunization. Fol. Primatol., 22: 59-71.
Moor-Jankowski, J., A. S . Wiener, W. W. Socha, E. B. Gordon and J. H. Davis 1973 A new taxonomic tool: Serological reactions in cross-immunized baboons. J. Med. Prim.,
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