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Blood groups as genetic markers in chimpanzees Their importance for the national chimpanzee breeding program.

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American Journal of Primatology 1:3- 13 (1981)
Blood Groups as Genetic Markers in Chimpanzees:
Their Importance for the National Chimpanzee
Breeding Program
w.w.
SOCHA
Primate Blood Groups Reference Lahorntory and World Health Organization Collaborating Centre
for Ha.emntology ofPrimate Animals and Lahoratory for Experimental Medicine and Surgery in
Primates (LEMSIPI of the New Yurk University School of Medicine
Severe restrictions on the importation of chimpanzees emphasize the importance and urgency of domestic breeding as a sole means to assure an
uninterrupted supply of animals for medical research. An insight into the
genetic structure of the self-sustained captive population of animals is indispensable to prevent the effects of inbreeding and to preserve the animals’
reproductive capacity. This can be achieved by study of sets of genetic markers
in the form of heritable molecular or antigenic variations detectable by relatively simple methods. Among chimpanzee blood components so far identified as
possible genetic markers, red cell antigens appear to be the most useful and most
readily available. The amount of information concerning blood groups of chimpanzees, their serology and genetics, number of polymorphic types, etc, surpasses data on other heritable traits in this species. A concise review of the
present status of knowledge of chimpanzee blood groups and, particularly, of
serology and genetics of two complex blood group systems, V-A-B-D and
R-C-E-F, is given together with a few examples of their application in cases of
disputed parentage. Finally, a list of practical steps is suggested dealing with
introduction and use of genetic markers as elements of the national chimpanzee
breeding program.
Key words: chimpanzee, blood groups, genetic markers
INTRODUCTION
The recently adopted Convention on International Trade in Endangered Species of Wild
Flora and Fauna restricts the movement of chimpanzees across international borders
and may ultimately ban their importation for research purposes. In view of continuous,
and probably increasing importance of chimpanzees in bioscientific programs, particularly in medical research on several major human diseases, steps must be taken to
assure an adequate supply of these invaluable animals.
Received October 15, 1980; accepted November 19, 1980.
Address reprint requests to Wladyslaw W. Socha, MD, New York University Medical Center, 550 First Avenue,
New York. NY 10016.
0275-2565/81/01014003$03.500 1981 Alan R. Liss, Inc.
4
Socha
A 1978 report of the Task Force on the Use and Need for Chimpanzees recommends,
among other things, that:
-Government support should be provided for the domestic breeding of enough
chimpanzees to meet our national bioscientifk requirements on a continuing, permanent basis;
-A national, coordinated long-term program should be designed and implemented
to maximize the breeding potential of chimpanzees already available in the United
States. . . .” [Report of the Task Force, 19781.
The specifics of the national chimpanzee breeding program were further elaborated by
an ad hoc panel of consultants who made a number of recommendations dealing with the
creation of a self-sustained captive population of chimpanzees which would satisfy
essential requirements ofthe bioscientific community. The report of the panel stated that
“special attention [should] be given to the genetic aspects of the captive chimpanzee
population” [Ad Hoc Consultation on Chimpanzee Breeding, 19791. The report on the
“Cemus and Inventory of the U.S. Chimpanzee” [Seal et al, 19801, prepared on the
recommendation of the ad hoc panel, identified two areas in which availability of genetic
data seemed to be of utmost importance for the future of the large-scale breeding
program: establishment of pedigrees of captivity-born animals and surveying changes
occurring in the gene pool of captive populations in order to prevent inbreeding and to
maintain genetic diversity indispensable for preserving optimum breeding capacity of a
population effectively cut off from supply of feral animals.
The real and imminent danger of progressive impoverishment of the genetic patrimony
of the self-sustained populations stems from known patterns of breeding practices, which
tend t o be built around a few, selected animals known for their exceptionally good
reproductive performance. This implies that practical means have to be found to gain
insight into the genetic structure of the collection of animals at our disposal and to
monitor the dynamics oftheir gene pool, both in time and in space. Obviously, it would be
impractical, and probably impossible, to identify and count each of the sections ofDNA on
the chromosomes of over a thousand chimpanzees. It is possible, however, to achieve the
goal by indirectly identifying and counting, by methods of population genetics, some of
the genes from the occurrence of their products, which are directly accessible to investigation.
Although one could list a number of heritable traits, both physical and biochemical,
only some of them meet requirements of genetic (or chromosomal) markers. By definition, the genetic markers are body components that exhibit heritable variation
sufficient to be classified as genetic polymorphisms. Polymorphic traits are only those
traits that occur in a population more frequently than would be expected on the basis of
recurrent mutations. For example, the so-called inborn errors of metabolism usually do
not satisfy this requirement. Among other characteristics that suggest the usefulness of
a given trait as chromosomal marker are 1)its discontinuous and unitary nature;
2) detectability in unchanged form, from early stages of life until death of an individual,
by relatively simple and easily reproducible techniques; 3) unequivocal mode of inheritance, possibly by individual genes, sharply differentiated from one another; 4)existence
of several polymorphic variations within one and the same locus; and 5) frequencies
varying significantly from one population to another.
The genetic markers most commonly used are either molecular or antigenic variants,
which are either immediate gene products or, at least, final products of relatively short
chains of reactions initiated by a gene, so that the chances of extra-hereditary factors
influencing the appearance of those markers are negligible.
Among numbers of such heritable variants, the components of blood have been the
objects of most extensive investigations. From the available literature, which lists over
40 molecular components of chimpanzee blood investigated, only ten have so far been
Blood Groups of Chimpanzees
5
TABLE I. Blood Components as Possible Genetic Markers in Chimpanzee
Number of
polymorphic
types
Authork)
I.
Plasma
Immunoglobulins (Gm, Inv)
Transferrins
Group-specific component
(Gc)
Alkaline phosphatase
11. Fkd cell isozymes
Glucose-6-phosphate dehydrogenase (G-6-PD)
Nicotinamide adenine dinucleotide phosphate
diaphorase (NADPH
diaphorase)
Peptidase A (triallelic)
Peptidase C (triallelic)
Phosphoglucomutase
(diallelic)
Phosphogluconate dehydmgenase (diallelic)
111. White cell antigens
Major histocompatibility
complex ChLA (A & B loci)
IV. Red cell antigens
V-A-B-D and R-C-E-F blood
group systems and sets
of unrelated specificities
2
8
2
Alepa [1968]
Goodman [19681
Cleve & Patutschnick [1979]
2
Lucotte [1980]
3
3
Beutler & West [19781
Lucotte 119801
Meera Khan & Balner I1972 I
4
4
3
Meera Khan & Balner 119721
Meera Khan & Balner 119721
Meera Khan & Balner L19721
3
Meera Khan & Balner 119721
18
Balner e t a1 [1978]
67
Socha & Moore-Jankowski [1979,
1980; unpublished data]
found polymorphic, thus meeting the basic requirement of genetic markers. Table I lists
those variants, together with two classes of heritable antigenic types of blood cells.
The choice of genetic markers for routine testing should be based on the relative
usefulness of a given set of markers, and the ready availability of the testing tools
(reagents, equipment, adequate controls, and-above all-experienced personnel).
The usefulness of a system of markers, which in the case of paternity investigations can
be mathematically expressed, is by and large, measured by the number of polymorphic
forms (complexity of the genetic structure) and depends on favorable distribution ofthose
types in the population [Wiener & Socha, 19761. As shown in Table I, the white and red
blood cell antigenic systems of chimpanzees offer the highest degree of polymorphism.
Although no data are yet available on the relative usefulness ofvarious genetic markers
of chimpanzees, such information recently became available on pig-tailed macaques
[Chakraborty e t al,19791. In that study, exclusion probabilities were estimated for each
of the 12 polymorphic red cell proteins and two independent red cell blood group systems,
the A-B-0 and Drhgraded systems. The latter two, which together offered eight phenotypes, contributed more than one-half of the cumulative probability of exclusion obtained
when all 14 systems were applied. It can be expected that in chimpanzees, the contribution of blood groups (with their 67 polymorphic variations) would be even more significant.
6
Socha
TABLE IT. Blood Groups of Chimpanzee: Present Status
Blood group
system
Macrospecificities
Microspecificities
Number of
phenotypes
A-B-O
0
A
AYh, A:;,,
M-N
M
v.0, v.A, v.B, v.D, v.AB,v.AD, v.DB, w
MN
or
MNV
V..o, V.A, V.B, V.D, V'.O, V'.A, Vq.B,
Vq.D, V"q.0, VDq.A,VPq.B, VPq.D,W
Rare forms: Vuu5.0,Vpq'.A, VPu'.B,
V""'.D
rh-negative
rc,, r q , rCF, rCFc, , rCFcL
Rh-positive
Rc,, Rc2, RC, RCc,, RCq, RCE, RCEc,,
RCEc,, RCF, RCFc,, RCFc,, RCEF, RCEFc,,
RCEFc, ,
Rare forms: Rdb,R"', RD', R, R,,,
24
Hc, he; G", g'; K', k (very rare);
0 , M ' , mc; Nc, n#';Tc, t c
14
Rh-Hr
Probably
unrelated
specificities
Aih
Total number of phenotypes defined
4
25
67
There are, however, several other factors that suggest the exceptional value of blood
groups as genetic markers of chimpanzees.
1)Studies of heritable antigenic properties of chimpanzee red cells were started as early
as 1925 ILandsteiner & Miller, 19251 and have been conducted systematically for the
last 20 years. The resulting amount of information on chimpanzee blood groups approaches the level of knowledge of the human blood groups.
2 ) Blood grouping techniques used for testing red cells of chimpanzees are identical, or
almost identical, to those routinely employed i n human blood grouping practice and
proved in millions of tests [Erskine & Socha, 19781. The results of blood grouping tests
in chimpanzees were found to be fully reproducible, and the specificity, titers, and avidity
of reagents available in our laboratory were tested on a large number of samples.
3) Modes of heredity of chimpanzee blood groups were established by population studies
and, on many occasions, were tested on chimpanzee families and harems.
4) In addition to their application as genetic markers, the blood groups have practical
value in the management of colonies of captive chimpanzees, and in some experimental
uses of those animals. The knowledge of blood groups is indispensible in clinical and
experimental situations calling for transfusion of blood or transplantation of organs and
tissues. Materno-fetal red cell incompatibility seems to be responsible for at least some
cases of transplacental immunization.
5 ) Finally, but not least important, blood group records are already available on over
350 chimpanzees: that is, according to the most recent census [Seal e t al, 19801, blood
group records are available for close to one-third of the total population of chimpanzees
currently maintained in the United States.
Blood Groups of Chimpanzees
7
TABLE 111. Serology and Genetics of the Chimpanzee V-A-B-DBlood Group System
Number
Reaction with serum of specificity
Designation
Anti-V“
Anti-AC
Anti-BC Anti-D‘
+
+
c
+
+
v.0
V.A.
V.B.
V.D.
+
+
u
=
ud
=
=
vR
lill
=
V
=
-
+
-
-
+
-
-
Possible
genotypes
Observed Expected
0
43
27
5
49
9
12
0.45
43.00
27.00
3.20
53.30
13.20
10.10
uu
17
47
27
9
17.0
40.6
31.2
7.7
VV or Vu
Vu”
VuH
VuL’
ir4ud or u 4 u
didi or uu
uDuu or iPu
uAuH
u4u”
uUuU
Estimated gene frequencies
0.0429
0.3761
0.2891
0.0715
0.2206
x’,~,= 5.310, 0.5 < P < 0.70
Table I1 summarizes the present status of knowledge of chimpanzee blood groups.
Technically, there are two levels at which the testing of chimpanzee red cells is approached and which stem from the principle of “immunological perspective” [Landsteiner, 19361:
1)With the help of agglutinating reagents originally developed for use on human red
cells, macrospecificities [Wiener & Socha, 19741on the red cell membrane are recognized
that probably are common to all Hominoidea, including man.
2) Further subdivision of antigenic types is accomplished with the use of narrowly
specific reagents produced either by cross- or isoimmunization of chimpanzees. Specificities detected by those antisera are, by and large, proper to chimpanzee red cells alone,
although some of them were also found on red cells of other species of Pongidae [Socha
& Moor-Jankowski, 19791.
The blood groups of the former class were previously described as “human-type,” a term
recently abandoned as misleading and, perhaps, too anthropocentric, since at least some
of the so-called human-type specificities probably existed well before the emergence of
man [ R e i e & Socha, 19801.
However, sharp separation of the second class of specificities, previously known as
“simian-type”blood groups, also seems unfounded. Specificities of that type are no longer
considered exclusive attributes of the red cells of nonhuman primates; they were also
detected, in polymorphic forms, in human red cells [Socha & Moor-Jankowski, 1978,
19791.
One of the two complex chimpanzee blood group systems so far defined, the V-A-B-D
system, is the counterpart and an extension of the human MNSs system. The so-called
R-C-E-F system is the chimpanzee counterpart and extension of the human Rh system.
8
Socha
TABLE IV. Serology and Genetics of the Chimpanzee R-C-E-F Blood Group System
Reaction with
serum of
specificity anti-
Number
Designation
R" Cc EC F' c' cf Observed Expected"
rc,
_ _ _ - + +
rc2
-
rCF
rCFc,
rCFc2
Rc,
- + - + - -
Rc,
+ - - - + -
RC
RCc,
RCc,
RCE
RCec,
RCEc,
RCF
RCFc,
RCFC,
RCEF
+ + - - - -
RCEFc,
RCEFc,
+ + - + + + + + + - -
53
3
2
18
5
37
16
0
2
2
2
6
4
11
19
16
6
50.17
2.84
2.17
16.52
4.97
05.02
15.15
0.02
1.64
1.18
0.80
9.05
6.94
8.13
19.87
13.97
5.60
+ + + + + +
+ + + + + -
1
0
1.32
0.95
-
.
.
-
+ -
- + - + + +
t - t + + - - - + t
-
+ + - - + +
t
+
-
-
+
-
+ + + - - + + + - + +
+ + + - t -
+ + - + - + + - + + +
Possible genotype(s)
x'(lo, = 7.6023
0.70
P
0.50
"Based on estimated gene frequencies:
=
=
0.39272
P
i-l
R'
= 0.02254
R'
=
0.05369
R l ~ E
R'~7' =
~
0.11828
0.17983
0.11225
rIF
=
R'
R'"'
=
0.10357
0.00974
0.00783
Serology and genetics of the V-A-B-D and R-C-E-F systems are presented in Tables 111
and IV respectively.
For detailed descriptions of both systems, the reader is referred to earlier articles
I Wiener et al, 1974; Socha & Moor-Jankowski, 1979, 19801.
To illustrate practical applications of blood groups in reconstruction of chimpanzee
pedigrees, three cases are presented (Tables V and VI) in which blood grouping testswere
instrumental in solving problems of dubious paternity.
CASE 1
Two males were taken into consideration a s possible fathers of "Tucson'' and
"Gretchen," two offspring of the female "Martha" (Table V, Ch-138). The question of
paternity could be solved by excluding "Oscar" as the father of Tucson, and at the same
time, by eliminating Walter as the father of Gretchen. Tucson was found to be group RCF,
a type that could not have come from its mother or from male Oscar, but could have been
inherited from the male "Walter." Walter, in turn, had to be excluded as the father of
Gretchen, who was of phenotype rc, , to which neither of Walter's R-C-E-F alleles could
have contributed. In addition, Gretchen was typed V.A, whereas both her mother and the
0'10'
T' IT'
Or
T'
Case 1
Rr lrr
H' IH
K' IK'
NLlnc
010
VPYIUR
MNIMn
0
MN
V""B
RCF
H"
K'
N'
WALTER
(Ch-168)
RLlr'
H'IH'
K'IK'
ncIn'
0' lo'
F IF
VIUU
A' lA2
MNIMN
0
T'
nL
n'ln'
Q lo'
T'lF
TUCSON
(Ch-285)
A2
A210
MN
M NIMN
V.B
VIVR
RCFc, RCFlr'
H'
HI IH'
K'
K' IK'
0'
t'
nc
A,
MN
VB
Re,
H
K'
MARTHA
(Ch-138)
K'IK'
n' In"
oc10'
T'IT'
H'IH'
A'IA'
MNIMN
Vlu"
RcLlrl
n' In'
0' 10'
T'IF
n'
OC
T'
GRETCHEN
(Ch-248)
A,
A'IA'
MN
MNIMN
V.A
VIUL
rc I
rllri
H"
IJ' lHf
K'
K' IK'
A'
MN
V.A
RCEc,
H'
K"
nc
0'
T'
OSCAR
(Ch-211)
T'
OC
nr
H'
K'
Rc,
A,
M
v.AB
R2P
H' IH'
K1IK'
n'ld
0
' lo'
T' Itc
u'Iu~
A'IA'
MnlMn
TABLETOP
(Ch-467)
K' IK'
n' In'
O'IOC
T'IT"
K'
Case 2
T"
0 c
nc
h' lh
rilr'
A'IA'
MnlMn
U ~ I L ~
h'
A,
M
v.A
rcl
OLGA
(Ch-22)
K'IK'
n"lne
oc lo'
T'lt'
R%f
H' IH'
V"IUR
Mnl Mn
All0
JUSTIN
(Ch-315)
A,
A'IA'
M
Mnl Mn
v.AB
u4liu"
R,Cc,
R$ h'
H'
H' lh'
K'
R /K'
N'
nLIn'
0'
0' lo'
T"
T'IT'
T'
OL
K'
n'
w
A,
M
v.B
&CF
RUFE
(Ch-114)
blood group phenotypes of animals involved, as well as their genotypes. The latter were inferred directly from pedigrees of larger
families andlor harems, not shown in this table, or were calculated as "most probable genotypes" from the gene frequencies.
TABLE V. Examples of the use of blood groups for solving problems of parentage in chimpanzees. For details, see the text. Shown are
10
Socha
TABLE VI. A n o t h e r Example of the Use of Blood Grouping Tests for Solving Problems of
Parentage in Chimpanzees (Case 3)*
Animal's identification
Putative sire I
Ch-2019, BAMRAM I
Putative sire I1
Ch-2003, KOBI
Putative sire 111
Ch-2004, GERONIMO
Blood type
A2
M
v.AB
rc1
0'
nln
t'
A,
M
v.D
RCE
A'IA' or AIIAz or All0
MnlMn
u"h" or v"lv
R"iR'" or R''lR'
tit
OC
010
NC
H'
N I N or N l n
HIH or Hlh
TIT or Tit
AIA or A10 (not tested for A, 1
MnlMn
@lv"
R" IR or R" Ir or Rlr" (not tested for cI I
(not tested for 0")
nln
HIH or Hlh
(not tested for TL)
A21A2or AZIO
M N I M N or M N ! M n
V!U
+ / F ~ or rll?
010 or 010
NIN
Hlh
TIT or Tlt
A'IA' or All0
M N I M N or M N I M n
Vlud
R'IR' or RIlrI or R'IR2 or R'IP
010 or 010
nln
Hlh
T'
A
M
A2
MN
V.A
3
0
n'
H'
T'
A,
MN
V.A
Rc,
0'
n'
H
t'
A>
MN
VO
Rci
0
n'
H
1v
"For details, see t e x t
r'lr' or r l l P
010 or 010
HIH or Hlh
n'
Offspring
Ch-1036, JAYME
UAluB
H'
H"
Female parent
Ch-1027, KISSEY
AIIA' or A2lO
MnlMn
n'
V.AD
RCFc
Putative sire IV
Ch-2020, COCOA
Possible genotype(s)
tit
ALIA2or A210
M N I M N or M N I M n
V!V or Vlu
R'IR' or R'IP or R'lr'
010 or 010
nln
HIH or IIIh
Tit
Blood Groups of Chimpanzees
11
male Walter were of type V.B. Since specificities rc, and V.A were parts of the phenotype
of the second male involved-namely, of Oscar-he had to be assigned as the father of
Gretchen.
CASE 2
The second case constitutes not only another example of exclusion of one of the two
putative fathers, but also a rare instance where a male could be positively identified as
the father. In this case, the male “Tabletop” (Table V, Ch-467) had to be excluded as the
father of the baby “Justin,” since neither of the male’s two R-C-E-F alleles iR2 or P) could
have contributed to the baby’s genotype. On the other hand, paternity ofthe second male,
“Rufe,” was positively demonstrated: both Rufe and baby Justin displayed a very rare
varient of specificty Rc, designated R%,found in only one male and one female of
age from among 410 chimpanzees so far tested by us.
CASE 3
Case 3 involved four males as putative sires of the baby chimpanzee “Jayme,” offspring
of the female “Kissey” (Ch-1027). As can be inferred from the data in Table VI, two ofthe
males, “Bambam I” and “Geromino,” had to be excluded on the basis of their V-A-B-D
groups since neither of them could have contributed to the type V.0 of the baby. The male
“Kobi,” on the other hand, was excluded as the father on the basis of his R-C-E-F type:
none of the alleles that made up his possible genotypes could have contributed to the
blood type of Jayme. Only the fourth of the males, “Cocoa,”was not excluded as the father
of Jayme. Unlike the situation in Case 2, however, the blood groups of Cocoa, as well as
those of the mother and of the baby, belong to types quite commonly found among
chimpanzees, and this precludes definitive identification of Cocoa as the father of Jayme.
Another field where blood groups of chimpanzees can possibly find practical application
is so-called sero-prirnatology-ie, study of distributions of blood groups in various geographically or otherwise separated populations for the purpose of identification or confirmation of subspecies. In a series of such studies, significant differences were found in
the distribution of some of the blood types among groups of chimpanzees previously
classified as separate subspecies on the basis of their morphological characteristics
[Moor-Jankowski and Wiener, 19721. The results indicate that blood groups could be a
part of the set of parameters used for taxonomic classification of common chimpanzees.
Blood grouping tests carried out on limited numbers of pygmy chimpanzees (Punpuniscus) showed them all to have blood types that are very rarely observed among common
chimpanzees [MoorJankowski et al, 19751. The occurrence of those rare blood types in a
chimpanzee of presumably common type may be indicative, even in the absence of
morphological evidence, of a not-too-distant hybridization with pygmy chimpanzees. It
must be stressed, however, that the reliability of blood groups as a tool for taxonomic
classification of chimpanzees is still questionable in view of the small size of samples so
far studied and lack of comparative data on feral animals.
In these early stages of preliminary plans for a national chimpanzee breeding program,
it may be timely to recall that about 40 years ago, the practical needs for optimization and
rationalization of breeding of domestic animals brought about a n explosive development
in the research of blood groups and other chromosomal markers in various species of
domestic and domesticated animals. Until now, immunogenetics in general, and blood
group serology in particular, of cattle, swine, horses, chickens, sheep, etc have developed
into separate disciplines practiced in laboratories all over the world, and there is a
prolific literature of the subject [for review of bibliography, see Irwin, 1956; Stormont,
1962; Andresen, 19631. Today, the blood typing is routinely used for identification of
individual animals in ascertaining the validity of registration and to solve problems of
parentage arising in the registration ofpurebred animals. Because of the widespread use
of artificial insemination in cattle and because of potential use of frozen semen years
after animals from which it is collected have died, the Purebred Cattle Association of
12
Swha
America ruledmany years ago that all registered dairy bulls used for artificial insemination must have their blood groups oficially recorded. Subsequently, dairy-breed organizations in many parts of the world adopted similar rulings and, as a consequence,
numerous laboratories for cattle blood typing have developed to meet the needs of the
industry.
The importance of blood grouping tests was later recognized for the exclusion of
freemartinism and monozygosity in twins and higher multiples, both in cattle and sheep,
Blood group loci of chickens are still used in measuring residual heterozygosity and
combining ability of heterosis.
Drawing from vast experience gained through use of immunogenetic data for practical
purposes of the animal breeding, one can outline, in general terms, steps to be taken to
make the best use ofthe available genetic markers in the upcoming national chimpanzee
breeding program INational Primate Plan, 19801:
1) Selection of a standard set of genetic markers on the basis of their usefulness and
availability.
2) Carrying out tests for standard markers in alE animals currently maintained in the
country, or, at least, in animals presumed to be candidates for the national breeding
program. Information obtained in this way should give adequate insight into the extent
of genetic diversity of the captive population and, at the same time, serve as the baseline
value for the future comparisons of the genetic census data. Computerization of the data
would be essential for the speedy storage, retrieval, and processing of accumulated data.
3 ) Continuous updating of the existing data by tests on all new acquisitions (import,
exchange, newborn animals).
4)Determination of geographic and/or subspecies variations by correlating data on
genetic markers with physical taxonomy classifications. If feasible, comparison of data
on captive animals with those obtained from field studies on representative samples of
feral chimpanzees.
5) Establishing genetic profiles of families and harems. In doubtful cases, reconstruction of pedigrees or paternity by means of available genetic markers.
6 ) Long-term genetic population survey by means of periodic gene frequency analyses
in entire population, in geographic sub-classes, breeding colonies, harems, and families,
to detect possible effects of inbreeding.
7) Ancillary studies of possible correlations between genetic markers and breeding
performance of individual animals, linkage analysis, etc. Accumulated data on all blood
groups of all animals, easily retrievable, may find practical clinical applications in cases
requiring blood transfusion, in experiments on transplantation of organs and tissues, as
well as in prevention of transplacental immunization due to materno-fetal incompatibility.
It is not certain whether the current technical means and existing facilities would allow
introduction of a multitude of various genetic markers for simultaneous testing in large
groups of animals. At present the red cell antigens of chimpanzees seem to be the prime
target, and the easiest one. The blood grouping of captive chimpanzees in the United
States has been continued for many years, and has already been accomplished for a
significant part of the total population of animals. The other markers such as red cell
isozymes of histocompatibility systems could be added at any moment, provided that
sound theoretical bases for their use are established and logisticproblems of testing large
numbers of animals are solved.
ACKNOWLEDGMENTS
The work was supported by USPHS, NIH grant 12074 and contracts RR 01082 and
RR-4-2184.
The author wishes to thank Mrs. J o Fritz, Executive Secretary, Primate Foundation of
Arizona, Tempe, Arizona, for providing blood samples of a chimpanzee family and for
permission to include the results of blood grouping tests in this report.
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13
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REFERENCES
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