Blood groups of bonnet macaques (Macaca radiata) with a brief introduction to seroprimatology.код для вставкиСкачать
Blood Groups of Bonnet Macaques (Macaca radiata), with a Brief Introduction to Seroprimatology W. W. SOCHA, J. MOOR-JANKOWSKI, A. S. WIENER, D. R. RISSER AND H. 'PLONSXI Primate Blood Group Reference Laboratory and W o r l d Health Organization Collaborating Center f o r Primate Hematology, Laboratory f o r Experimental Medicine and Surgery in Primates ( L E M S I P ) o f t h e N e w York University School of Medicine, and t h e Department of Anthropology, University of California, Davis, California KEY W O R D S Macaques . Bonnet macaques . Crab-eating macaques . Primates . Blood groups . Seroprimatology. ABSTRACT The human-type A-B-0 blood groups of 52 bonnet macaques ( M a c a c a radiata) were determined. Application of method of population genetics indicated the gene frequences to be 0 = 0.173, A = 0.480 and B = 0.347. Cross testing of sera and red cells of the bonnet macaques revealed two bloodtype-specific isoagglutinins, one of them strong enough for use as a blood typing reagent. No blood group polymorphism was revealed by testing bonnet macaque red cells with isoantisera produced in rhesus monkeys ( M . mulatta) and in crabeating macaques ( M . fascicularis). The rhesus and crab-eating macaque isoantisera reacted either with all or with none of the bonnet macaque red cells tested. The present report is the continuation of our studies on what we have defined as the human-type (Wiener, Moor-Jankowski, Gordon and Shell, '66), and simian-type blood groups of apes and monkeys (MoorJankowski and Wiener, '65). This work began with our early discovery of human blood factor M in the rhesus monkey (Landsteiner and Wiener, '37) and has resulted in about 150 publications providing an over-view of the presence and distribution of both human-type and simian-type blood groups in various species of apes and monkeys, most recently summarized in 1972 (Moor-Jankowski and Wiener). The importance of this body of information for anthropology lies mainly in the recognition of the presence in simians of blood groups homologous (Wiener et al., '72), i.e., human-type, and analogous, i.e., simian-type, to the human blood groups. In the course of our work, it became apparent that some of the simian-type blood groups detected by the isoimmune sera prepared by us in one species had homologues in other primate species in which these antisera detected polymorphic blood group kM. I. PHYS. AhTHROP., 45: 485-492. differences, viz, chimpanzee C-E-F system in orangutans and gorillas (Moor-Jankowski et al., '73b) and rhesus monkey isoantisera showing polymorphisms in crabeating macaques (Moor-Jankowski et al., '67). On the other hand, there also exist isoantisera produced in one species which react with all or none of the red cells of other primate species tested as shown in the present article. The observed cross reactions of blood grouping sera with red cells of various primate species have been developed into a new taxonomic approach to the study of nonhuman primates, as outlined in two recent papers (Moor-Jankowski et al., '73a; Moor-Jankowski et al., '74). Moreover, already in 1964 (Moor-Jankowski and Wiener, '65) we observed statistically significant differences between the distributions of blood groups in two population groups of baboons of different origins. Further studies have enabled us to extend this observation to differences i n blood group distribution among populations of chimpanzees (Moor-Jankowski et al., '66), to larger series of baboons (Wiener and Moor- 485 486 SOCHA, MOOR-JANKOWSKI, WIENER, RISSER AND PLONSKI TABLE 1 Results of A-B-H grouping of a series of 52 bonnet macaques (Macaca radiata) Reactions given by Serum 1 Saliva Typical results A and H B and H A, B and H Atypical results a A and H A, B and H A, B and H A, B and H Incompletely tested Not done Not done Not done Presumed blood group Number of animals Anti-B Anti-A None A B AB 18 12 15 Anti-B, weak anti-A Anti-A, anti-B Anti-B Weak anti-A A AB AB AB 1 1 Anti-B None Anti-B and weak A AB A 1 1 1 1 1 anti-A, ~ 1 As determined after absorption with human group 0 red cells. (Before absorption titers for human group 0 red cells ranged from less than 1 up to 8 units by saline agglutination method.) 2 Not conforming with Landsteiner's rule. The saliva samples were tested by the quantitative inhibition technique for the presence of the A-B-H blood group substances. The serum samples were tested for anti-A and anti-B agglutinins after preliminary absorption with human group 0 red cells. The red cells were tested for simian-type agglutinogens by using our isoand cross-immune sera prepared in rhesus monkeys and in stump-tailed macaques (Wiener et al., '73). The red cells and sera of the bonnet macaques were crossmatched against one another. All the tests performed in our Primate Blood Grouping Reference Laboratory were carried out by the same methods as used MATERIALS AND METHODS for typing of human beings for clinical and All the animals studied were maintained forensic purposes, which have to guaranby the California Primate Research Center tee precise, reproducible results. Thus, only at the University of California, Davis, Cali- specific reagents were used, the high pofornia. Eight of the animals tested were tency of which was confirmed by titration wild-caught, while all the remaining ani- against control red cells of known types mals were born since establishment of the routinely included in every experiment. DeColony at the Primate Center. The animals tails of methodology have been described tested therefore did not constitute a pan- elsewhere (Socha et al., '72a; Erskine, '73). mictic population. The blood and saliva RESULTS samples were collected under the supervision of one of us (D.R.R.) and shipped by The results of tests for human-type air mail to the Primate Blood Group Refer- A-B-0 blood groups are shown in table 1. ence Laboratory at the Laboratory for Ex- With only relatively few exceptions, Landperimental Medicine and Surgery in Pri- steiner's modified rule holds i n bonnet mamates, New York. The saliva specimens caques, namely, those and only those aghad been inactivated by boiling for twenty glutinins (anti-A or anti-B) are present minutes immediately after collection, be- in the serum for which the corresponding fore being shipped. group-specific substances are absent from Jankowski, '69) and to gibbons (MoorJankowski and Wiener, '67), thus estabishing a new discipline of seroprimatology (Moor-Jankowski and Wiener, '67), analogous to the discipline of seroanthropology, viz the study of the distribution of blood groups in various human populations, as initiated by the Hirszfelds (Hirszfeld and Hirszfeld, '19), and expanded in modern times by Mourant ('75) and by Ruffik (Bernard and Ruffik, '66). The present paper reports on the results of the extension of our blood grouping tests to a series of bonnet macaques (Macaca radiata). 487 BLOOD GROUPS OF BONNET MACAQUES (MACACA RADIATA) TABLE 2 Results of cross absorption experiments with red cells of bonnet and rhesus macaques using antisera prepared in. rhesus monkeys and crab-eating macaques Antisera Anti-Grli (Rh-216) Anti-Crli ( Rh-4 ) Anti-Brh (Rh-2) Java monkey (J-1) ( anti-rhesus ) Titers for M . radiata M . mulatta 64 M . radiata M . mulatta Absorbed with red cells of M. radiata Absorbed with red cells of M. mulatta 0 0 0 256 64 4 0 4 0 M . radiata M . mulatta 12 6 0 0 0 0 M . radiata 16 128 M . mulatta 1 Unabsorbed 10 2 0 0 128 4 Species specific. the saliva. As in other species of Old World monkeys (Moor-Jankowski and Wiener, '72), the red cells of bonnet macaques failed to agglutinate with anti-A or a n t i 3 or anti-H reagents. The human-type A-B-0 blood groups were readily determined for the great majority of the animals; 18 monkeys were clearly group A, 12 were group B and 15 were group AB, but no group 0 animals were encountered. In four animals, the sera contained agglutinins in addition to those expected by Landsteiner's rule, as, for instance, a monkey with A and H in the saliva and weak anti-A as well as anti-B in the serum (see table 1, atypical results). These additional agglutinins did not react as auto-antibodies in bonnet macaques, but solely against human group A red cells. A similar situation had been encountered by us in baboons (Moor-Jankowski et al., '65) and is explained by the different chemical nature of the group A substance on red cells and in secretions. Therefore, the blood groups of these animals were determined on the basis of the blood group substances present in their saliva, and confirmed by the strong reciprocal agglutinins present in their sera, while weak additional agglutinins were discounted. Sera but not saliva had been received for three of the monkeys; the A-B-0 blood groups of those monkeys were inferred by applying Landsteiner's rule and discounting any very weak agglutination reactions. The gene frequency analysis was based solely on the results for 45 animals for which complete, entirely typical reactions were obtained. From the observed fre- quencies of blood groups: 0 = 0.0, A 5 0.4000, B = 0.2667, AB = 0.333, by using the method of direct count, the gene frequencies prove to be A = 0.5667 and B = 0.4333, if one assumes gene 0 to be absent from this species. If from these estimated gene frequencies one calculates the expected frequencies of the blood groups and tests for goodness of fit, one obtains xZtr,= 4.64 so that P < 0.05. The fit between expected and observed blood group frequencies is much closer if one postulates the existence of gene 0 in bonnet macaques.' I n that case, the estimated gene frequencies using formulae of Wiener and Moor-Jankowski ('69) become 0 = 0.173, A = 0.480, B = 0.347 and test of goodness of fit gives x z c l , = 1.47 so that P > 0.20. By using the method of maximum likelihood, one of the referees who kindly reviewed this article obtained the following estimates of the gene frequencies: 0 = 0.1067, A = 0.5075 and B = 0.3859; and the chi-square resulting is 1.29 which is not much less than the value of 1.47 resulting when the gene frequencies are estimated by the use of our own formulae. Tests for simian-type blood agglutinogens were carried out with blood typing sera prepared in rhesus monkeys and crab-eating macaques (Wiener et al., '73). Positive reactions were obtained with a number of these reagents indicating the 1 The calculations presented here have the limitation that the population studied i s not a panmictic one. 685 MRA-379 390 399 430 442 481 482 588 596 645 657 Rh-2 Rh-3 Rh-6 Negative control ( M . mulatta) Rhesus monkeys 11. 12. 9. 10. 7. 8. 6. 1. 2. 3. 4. 5. ( M . radiata) Bonnet macaques Ked cells or - +++ +++ ++ ++ - ++ ++ ++ - ++ ++ ++ +++ + ++ ++ ++ ++ ++ ++ ++ (Rh-4) (Rh-3) (Rh-2) +++ +++ +++ +++ +++ -+it -t+ + +++ +++ +++ +i+ +++ A n ti-Crh Anti-Frh Ant i-Brh ++++ - - - - (Rh-6) Anti-Drh Reactions with antiserum ++ - ++ I I - - - - (Rh-212) Anti-Erh 3-+ - ++ ++ +++ +++ +++ +++ +++ +++ +++ +++ ++++ ++ +++ +++ (Rh-2 16) Anti-Grh Reactions ( b y antiglobulin method) of rhesus monkey blood typing sera with red cells of bonnet macaques including serum of one bonnet macaque (MRA-8) having a natural isoantibody TABLE 3 5 F 12 E ” 3F 2 E M 8 5z 2 +d +++ -t4--t +++ $ - :: +++ - ++ + - - MRA-8 ld 0 3: B * 5 v, BLOOD GROUPS OF BONNET MACAQUES (MACACA RADIATA) presence on red cells of bonnet macaques of agglutinogens homologous to those present in rhesus monkeys. That the reactions were indeed type-specific and not due to non-specific heteroagglutinins, was established by absorption experiments since, as shown in table 2, the antibodies anti-Brh, anti-@ and anti-Grh could be absorbed by positively reacting red cells from either bonnet macaques or rhesus monkeys. Of interest was the reaction of a serum from crab-eating macaques (J-l),immunized with the red cells of a rhesus monkey. This antiserum cross-reacted, although to a lower titer, with the red cells of bonnet macaques. As shown in table 2, antiserum J-1 could be fractionated by absorption with red cells of M . radiata indicating the presence in the serum of antibodies of at least two specificities, one shared by both M. mulatta and M . radiata and the other peculiar exclusively to M . mulatta. The red cells of the bonnet macaques were tested with the rhesus blood typing antisera in order to determine whether the latter could be used also for demonstrating individual differences in the blood of bonnet macaques. As shown in table 3, no polymorphism could be demonstrated since the rhesus reagents either agglutinated the red cells of all the bonnet macaques or none of them. When the sera and red cells of the bonnet macaques were cross tested no reactions occurred except in four cases, in which weak agglutination reactions occurred with the sera from some of the animals. In general, the specificities of these weak agglutinations differed from one another and the reactions were poorly reproducible. The clearest reactions were given by the serum of the female MRA-8 (cf. table 3 ) ; though weak, the reactions were distinct by the antiglobulin method. The findings were reproducible, though with difficulty, so that the serum could be used for reliable typing of bonnet macaques, provided that the tests were read blind and done more than once. In order to determine the presence or absence in the sera of heteroagglutinins for the human red cells, a series of unabsorbed sera from bonnet macaques were tested against human group 0 red cells by the saline agglutination methods. I n every case some agglutination occurred, though 489 in some the agglutination was very weak while in other cases the agglutination was quite avid, with titers ranging up to eight units. DISCUSSION The present investigation extends to another species of Old World monkeys our observations on human-type and simian-type blood groups of non-human primates, reviewed in the introduction to this paper. As in other Old World monkey species, the red cells of bonnet macaques do not react with anti-A, anti-B or anti-H reagents; nevertheless, their human-type A-B-0 blood groups could be determined by inhibition tests on the saliva and by tests on the sera for anti-A and anti-B agglutinins. The distribution of the human-type A-B-0 blood groups in bonnet macaques is similar to that observed by us for crab-eating macaques (Wiener et al., '74) but different from that of rhesus monkeys, all of which we have found to react as group B. While many of the blood typing reagents produced in rhesus monkey cross react with red cells of bonnet macaques, they proved to be of no value for blood typing the latter because those isoimmune rhesus sera which agglutinated bonnet macaque red cells reacted with red cells of all bonnet macaques tested, and were therefore useless for demonstrating polymorphism. These results show that when testing across specific lines, there is no essential difference between agglutinogens responsible for species-specific differences and those which give rise to type-specific diff erences. * For example, hum an-type blood group B may be considered a species-specific characteristic of rhesus monkey, but it is a type-specific characteristic in crabeating and bonnet macaques as well as in man. In contrast, simian-type blood factors, type-specific in rhesus monkeys, are apparently species-specific characteristics of bonnet macaque red cells. That typespecific differences ( polymorphisms) exist among the red cells of bonnet macaques was demonstrated by naturally occurring isoagglutinins found in four animals. Of these MRA-8, whose serum gave the most distinct reactions, proved to be a female of 2For a thorough discussion and definition of the concept of immunological species specificity as introduced first by Uhlenhuth ('01). the readers are referred to the paper of Moor-Jankowski et al. ('73a). 490 SOCHA, MOOR-JANKOWSKI, WIENER, RISSER AND PLONSKI breeding age. It is significant that she had had seven pregnancies, in three of which the infants died within a few months of birth. This supports the idea that the antibodies resulted from feto-maternal isoimmunization, as observed in marmosets (Gengozian et al., ’66) and in baboons (Volkova et al., ’66), and more recently in rhesus monkeys (Sullivan et al., ’72). For a detailed review and discussion of materno-fetal isoimmunization in primate animals the readers are referred to our recent paper on the subject (Wiener et al., ’75). In our studies we have not used the rhesus monkey antisera described on the one hand by Edwards and on the other hand by Duggleby and Stone (Stone, ’73; Edwards, ’73). Both groups used antisera diluted to one-unit titer while our work is directed by the standards required for human blood grouping. Edwards’ reagents were not available to us, while those of Duggleby and Stone were received in only small aliquots. Most of them gave nonreproducible results in our hands and the others detected the same specificities as our earlier described standard reagents. ACKNOWLEDGMENTS This study was supported by U. S. Public Health Service N.I.H. Grant GM-12074, Contract RR-4 12184. LITERATURE CITED Bernard, J., and J. Ruffie 1966 HBmatologie geographique. Masson et Cie, Paris (Vie). Edwards, R. H. 1973 Letter to the editor. Vox Sang., 24: 561-562. Erskine, A. G. 1973 The Principles and Practice of Blood Grouping. C. V. Mosby, St. Louis. Gengozian, N., C C. Lashbaugh, G. L. Humason and R. M. 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