TissueAntigens (1980),16,368-376

HLA Antigens in South American lndians

Francis L. Blackl, Lee Lucas Berman1 and Yvone Gabbay2 1Department of Epidemiology and Public Health, Yale Vniversity School of Medicine, New Haven, CT, V.S.A. and 2Instituto Evandro Chagas,Fundação Servicos de Saude Publica, Belém, Pará, Brasil

New HLA data for the Tirio, Parakanâ, Kayapo and Mapuche tribes, as well as supplementary data for the Waiâpi are presented. Taken together with previously published information on South American lndians, these typings show a remarkably homogeneous gene pool with a restricted range of polymorphisms and a further restricted set of haplotypes.

Receivedfor publication 14 january, revised,accepted 29 May 1980

In this report we present data on histo- Carib languageand their culture has charac- compatibility types from eight endogamous teristics commonly associated with the populations of five cultural and four Caribs. The capture of women from neigh- linguistic groups living in Brasil and Chile. boring tribes has been an established The number of specimens tested is small in practice. some instances, but when this is so a sub- stantial part of the endogamous unit is Kaxuyana: The Kaxuyana speak another represented. Coupled with previously pub- Carib language. According to Frikel (1970) lished data, these figures offer a consistent they are the survivors of several severely and reasonably comprehensive picture of depopulated tribes of the lower Trobetas the polymorphisms at the first three histo- River. Since 1970 they have lived adjacent compatibility loci in South American to the Tiriyo and roere has been some Indians. intermarriage.

Waiãpi (, Oyampi, Oiampi, Material Wajapi): Seventeen individuaIs have been Populations included in our study, comprising most of the population of the village of Molokopote Tiriyo (Trio): This group comprises S9 (1°2S'N, S3°S2'W). This population has related individuals from one village, Tirios, intermarried with the population of Trois near the Brasil-Surinam border (1°S7'N, Sauts in French Guiana previously studied SSo49'W) (Fig. 1) (Frikel 1961, Riviere by Tchen et aI. (1978). The language is of 1969, Black et aI. 1978). The Tiriyo use a the Tupi-Guarani group (Olson 1978) but

0001-2815/80/010368-09 $02.50/0@ 1980 Munksgaard, Copenhagen

~rial 369 HLA IN S. AMERINDIANS

social patterns and artifactual styles which are distinct froro the Waiãpi. The Parakanã Novo village raided the Assurini during the 1950s and now includes individuals of that lineage. The larger Velho village seeros to have been re!atively passiveand includes no known captives froro other tribes.

Kayapo (Cayapo): Our sample is drawn from three villages: the Xikrin from the northeast limit of the Kayapo range ° I o I . (6 30 S; 51 O W); the GorotIre a I arge group near the center of the range (7°20'S, 51°10'W); and the Mekranoti from the southwesterly edge of the Kayapo territory (8°39'S, 54°13'W). The three villages have 17 similar cultural patterns and share a Ge language. They probably separated by fission of a common village more than a c.entury ago (Vidal 1977), and within the memory of living individuaIs contacts have been entirely hostile. The Kayapo tradition- Figure 1. Map of showing loca- ally take captives, especiaIly children, from tions of populations discussedin this paper. other tribes and assimilate these persons 1. Vupa 10.Parakanã Novo into full tribaI membership. 2. Bari 11.Parakanã Velho 3. Warao 12.Xikrin Mapuche (Araucanians): We have included 4. Makiritare 13. Gorotire } Kayapo 87 unrelated individuals from Indian popu- S. Vanomama 14. Mekranoti lations of about 4,000 persons living in 6. Emerillon 1S.Quechua 7. Waiapi 16.Aymara several rural districts on the upper Biobio 8. Tiriyo watershed in Malleco province of Chile 9. Kaxuyana} C an .b 17.Mapuche18. (approx. 38o S 71 o W) Faron 1968, Etche- verry et aI. 1967, Black et alo1977). There many artifacts are similar in style to the is uncertainty as to whether the inhabitants neighboring Carib groups. of this alpine area represent chief1y descen- Parakanã (Parakanan): Our stUdy includes dants of the Pewenche,a relatively primitive most of the residents of two socially branch of the Mapuche who inhabited the separate but cultural1y similar villages area at the time of first contact, or refugees (locations at first contact: 4°S SI°30'W; from wars against the Spanish in the central 4"30'S SO030'W) (MagalhãesSantos 1976, valley of Chile. Although the area chosen is Black et aI. 1980). These villages also use a one of the most remote inhabited by the Tupi language but retain a number of Mapuche roere has been extensive accultUr- cultural characteristics ftrst noted in the ation and many Caucasian captives were extinct Tupinambaof the coastand utilize taken by the tribe during the 19th Century. r~'"''}--'\~ 370 BLACK ET AL.

Table 1 to confirm pattemspatterns of A19, A23-24 and Reaction pattems used to identify types 85, 15 B38-39 splits observed in the standard and w35 panel. Serum NIH R. Resu/ts N N eactlon ame o. The estimated gene frequencies for each Kraay 949 WK -- village are presented in Table 2. Gene Schmidt 774 + -- frequency was calculated by summation of VA260 625 + -- Murray 687 + -- haplotypes determined as described below Fe28-9 463 + -- in aU tribes except the Mapuche. For the Voort-Mol 1443 + NT- latter we used the formula f = 1 -~ Weismeyer 1017 -WK- where f = gene frequency and P = pheno- Templer 891 -+ - type frequency. The range of polymorph- AlIen 842 -+ + isms is very restricted except in the CC68 324 -+ WK Behard 1085 + WK + Mapuche where we found low frequencies Smith, D 779 --WK for several alieles which occur at higher Mol-Tuip 957 + WK + frequencies in Caucasians. The frequency Type Designation B5 B15 Bw35 of unidentified types was low, even for the C locus, for which only a narrow range of WK = Weak Reaction; NT = Insufficient Tests. serum specificities was available. The BS1- S2 split is not usualiy identified in the NIH Methods serum specificities, but one serum recorded Heparinized blood samples were trans- as BSl, V A260, was used in ali tests, and a ported to a laboratory at ambient tempera- second, Voort-Mol, was added for tests rufe. Lymphocyte separation was com- with the Tiriyo and Kaxuyana. AlI speci- pleted as described previously (Black et mens recorded as BS reacted with these aI. 1980) within 54h of sample collection. sera except two Waiãpi and one Tiriyo. Cytotoxicity tests were usually carried out Each of these was associated with Cwl. with a panel of 84 sera plus controls. For Only Bw39 of the B16 split was found in the most part, these were sera selected Brasil or in the Mapuche, but Bw38 has from the NIH bank to emphasize the types been reported in the Yupa (J ohnson et alo most often encountered in South American 1978). lndians. ln addition, we included sera with Also shown in Table 2 are unweighted Bw46 and Bw48 specificities contributed mean values of gene frequencies for 14 by Drs. Rose Payne, M. J. Simons and tribes. These tribes are: those listed here, F. Kissmeyer-Nielsen. This panel included with the two Parakanã, two Carib and two or more sera reactive for each of 13 A, three Kayapo viliages consolidated, and 18 B and four C alleles. Twelve or 13 sera with the Waiãpi data consolidated with with specificities for the B5, 15 w35 com- that of Tchen et alo (1978); and the plex were included. The reaction patterns Aymara (van der Does et alo 1973), the used to distinguish antigens of this com- Warao, Yanomama and Makiritare (Layrisse plex are shown in Table 1. The full seventh et aI. 1976); the Bari and Yupa (Johnson et workshop series of 176 antisera was used alo 1978) and the Emerilion (Tchen et aI. by Dr. Zulay Layrisse with 38 Parakanã 1978) and the Ticuna (Neel et aI. 1980). specimens,and these results have been used The spectrum of types encountered is HLA IN S. AMERINDIANS 371

Table 2 HLA gene frequency in tribes newly studied and mean afpublisbed data an 14 tribes No.Tested 73 531 98 116 87 11592 , '" Q o ~ ., "~ ..o. " .o .o ~ ~ ..=' "C .~ ".., ..>. o. .. ~ ~ ~ :t: ~ ~

A 1 O O O O 0.047 0.005 2 0.439 0.688 0.234 0.479 0.250 0.428 3 O O O O O 0.003 11 O O O O 0.006 0.001 w24 0.164 0.162 0.072 0.091 0.122 0.212 w26 O O O O 0.006 0.001 28 0.109 0.057 0.245 0.069 0.366 0.105 w30 0.007 O O O O 0.004 w31 0.278 0.020 0.438 0.360 0.059 0.0273 w32 O 0.066 0.005 O 0.090 0.011 w33 O O O O O 0.007 Blank O O 0.005 O 0.054 0.017

B 5 0.082 0.280 O 0.039 0.065 0.164 7 O O O O 0.012 0.004 8 O O O O 0.023 0.003 12 0.007 O O O 0.017 0.004 14 O O O O 0.078 0.007 15 0.007 0.007 0.019 0.319 0.142 0.133 16 0.205 0.330 0.208 0.134 0.156 0.176 17 O O 0.005 O O 0.002 18 O O O O O 0.002 w21 O O O O 0.017 0.003 w22 O O O O O 0.001 27 O O O O 0.017 0.003 w35 0.534 0.319 0.523 0.375 0.198 0.234 40 0.164 0.066 0.244 0.134 0.072 0.144 Blank O O 0.005 O 0.203 0.116 C wl O 0.088 O O O 0.0364 w2 O O 0.005 O O 0.001 w3 0.260 0.353 0.306 0.431 0.157 0.318 w4 0.527 0.382 0.478 0.409 0.293 0.348 Blank 0.212 0.176 0.211 0.160 0.550 0.296 1 Combined data from Tchen and present reporto 2 Unweighted mean of the tribes appearing in Table plus the Quechua (Tittor et ai. 1973), Aymara (van der Does et ai. 1973), Warao, Yanomama and Makiritare (Layrisse et ai. 1976), Emerillon (Tchen et ai. 1978) and Bari and Yupa (Johnson et ai. 1978), Ticuna (Neel et ai. 1980). 3 Undifferentiated Aw19 reactions reported for the Ticuna are incorporated into the Aw31 value. 4 C locus means based on six tribes only. 372 BLACK ET AL. remarkably consistent throughout the w39; and Cw1, Cw3 and w4, recur with continent. remarkable consistency in Indians from alI Most probable haplotype distributions tested South American tribes. This limited are given in Table 3. Where possible, these heterogeneity is further circumscribed by were determined Eram genealogies. When the fact that it includes the closely related incomplete genealogy or parental exclusions components of the A19 complex (except left uncertainty, haplotypes which had Aw29) and the 85, 15, w35 group, while been identified in the same village were excluding alI components of the A1, 3, assigned if possible. When more than one 10, 11 and 87,8,22,27 groups. (Amos & previously identified alternative presented, Ward 1975). the one with the higher frequency was Northern Amerindian populations are assigned. Thirty~even haplotypes were not generally asfree of Caucasianadmixture postulated, two of which, A28, B12, C- as the South American forest tribes and and -, B17, Cw2 were found in children most of the work on the northern popula- whose fathers are believed to have been tions was done before the HLA antigens Bush Negro and neo-Brasilian respectively. were as well-resolved as now. These factors On the basis of number of tests and gene make the identification of endogenous frequency, 77 haplotypes might have been types on the northern continent more expected if roere had been no disequi- difficult. In general, however, all the anti- librium. The same haplotypes were found gens which we have found in South America repeatedly in geographically distant tribes as well as types 8w21 and 827 occur in and one, Aw31, Bw35, Cw4, accounted for North American Indians in higher fre- 23% of alI sequences in the Indians of quencies than in Europeans (Cann et aI. lower Amazonia. 1973, 1974, Perkins et aI. 1974, Troup et The same methods for linkage analysis aI. 1974, Spees et alo 1974, Corley et alo could not be applied to the Mapuche 1974, Grofton et alo 1975). AIso, 814 sample of unrelated individuaIs, but stan- occ;urred at higher than European frequency dard disequilibrium analysis (Yasuda, 1978) in the Maya and possibly certain other shows high frequencies for the haplotype tribes. The Eskimo pattern, toa, is similar A2, Bw39, C- (D = 0.061; P by X2 = to what we have found, but they mar lack < 0.001) and A28, B5, Cw4 (D = 0.036; the A19 complex antigens while possessing P = < 0.001). The haplotype Aw31, Bw35, 8w22, 87 and the distinctive 8w48 antigens. Cw4, was not in significant disequilibrium (Kissmeyer-Nielsen et aI. 1974, Dossetor in this population. A1, 11 and w26, types et aI. 1974). Thus the South American not found in the northern tribes, were in tribes possessa subset of alicies which mar positive disequilibrium with B locus have been drawn from the slightly larger antigens 7, 8, 12 w21 and 27, likewise North American set. No other source need foreign to the northerners (p = 0.01). be postulated. On other continents, only However, B14, also absent in the Brasilian the Australian Aborigine exhibits similarly tribes, did not show this association. limited polymorphism over a comparable geographic arca (8ashir et aI. 1973). Discussion Within South America, only the presence The same sets of antigens, A2, 28, w24, of Cw1 in the north and low positive fre- w30, w31 and w32; B5, 15,40, w35 and quencies of Aw33 in the west and 814 in HLA IN S. AMERINDlANS 373

Table 3 Distribution of baplotypes in Indians of Para State, Brasil; frequency X 1,000 No. Tested 67 31 55 31 30 59 14 17 304

.;; ~ ,~ I~ O.. ~ .'-~ o 5 o c.~ ~.~ Village ~ "> ~ ~ ...o ~ ~ -.:4 O .~ ..o ;.. ;; ,~ , ~ ..~ Z o .:4 ...~ ~ .~ - ..> o ..~ ~ ~ ~ .~ ~ .~ ~ < Haplotype ~ ~ >< ~,.. ~

A2,B5,C- o o 18 O 117 25 215 O 30 A2,B15,Cw3 O O 27 65 100 O 36 O 23 A2,B40,Cw1 O O O O O O O 29 2 A2,B40,Cw3 127 O 36 O 17 25 O 29 43 A2,B40,Cw4 O O 55 97 O 42 O 29 30 A2,Bw35,Cw3 O O O 16 33 85 O 29 23 A2,Bw35,Cw4 15 97 191 129 217 212 36 294 142 A2,Bw39,Cw1 O O 9 O O O O O 2 A2,Bw39,Cw4 O O O O 50 8 107 O 12 A2,Bw39,C- 157 O 64 226 33 51 36 g 84 A28,B5,C- O O O O O O 36 O 2 A28,B12,C- O O O O O 8 O O 2 A28,B15,Cw3 O 32 36 O O O O O 10 A28,B40,Cw3 201 O O 113 O 51 36 O 67 A28,B40,Cw4 O O 27 O O 17 O O 8 A28,Bw35,Cw3 O O O O O 17 O O 3 A28,Bw35,Cw4 O O O O O 8 O 29 3 A28,Bw39,C- O 307 9 16 O 17 O O 38 Aw24,B15,Cw3 O 32 91 16 33 O O O 25 Aw24,B40,Cw3 O 48 O 16 17 8 72 117 20 Aw24,Bw35,Cw3 45 48 27 O 33 O O 176 33 Aw24,Bw35,Cw4 O O O O O 77 72 O 18 Aw24,Bw39,Cw4 O O O O O 59 O O 11 Aw24,Bw39,C- O O O O 17 17 O O 5 Aw30,Bw35,Cw4 O O O O O 8 O O 2 Aw31,B5,Cw1 O O O O 17 8 36 59 8 AW31,B15,C~3 O O 245 194 83 O O O 72 Aw31,B40,Cw3 O O O O 17 42 O O 10 Aw31,B40,Cw4 O O O O 17 O O O 2 Aw31,Bw35,Cw3 O O O O O 51 36 O 12 Aw31,Bw35,Cw4 447 419 164 113 183 128 143 O 231 Aw31,Bw39,Cw4 O O O O 17 8 O O 3 Aw31,Bw39,C- O O O O O 25 143 O 11 Aw32,B5,Cw4 O O O O. O O O 29 2 Aw32,Bw39,C- O O O O O O O 176 10 Aw32,B-,C- O 16 O O O O O O 2 A-,B17,Cw2 7 O O O O O O O 2 No. Homozygous 15 6 7 4 2 2 1 O 37

Homozygous Expected 19.0 8.9 7.9 4.5 3.6 5.5 1.7 2.9 54.0 BLACK ET AL.

the south suggest possible geographic discussedelsewhere (Black et aI. 1980). clines. In possessionof Cw1 and in haplo- In their study of the Waiãpi and Emeril- type frequencies, the Waiãpi are more lon, Tchen et alo reported that a common similar to other tribes north of the Amazon haplotype was A2, B blank. They did not which speak unrelated languages than to include B16 or Bw39 sera in their panel. In the Tupi speaking Parakanã who live 800 the light of our findings we believe that the kilometersto the south. haplotype they identified was most prob- Low levels ofcharacteristically Caucasian ably A2, Bw39, C blank. This same haplo- genes in the Mapuche, like those found by type was found in alI the Brasilian tribes, as Tittor et aI. (1973) in the Quechua, suggest well as more than 5,000 Km to the south post-Columbian admixture. The proportion in the Chilean Mapuche. of apparently exogenous alleles found in Bodmer (1975) suggested that the the Mapuche suggest 12% Caucasian ad- relative HLA homogeneity of American mixture when compared to data for Madrid Indians is due to "selective pressuresassoci- (Ryder et alo 1978) and agrees with ABO ated with infectious diseases", pressures tests carried out on the same population by which reduced HLA heterogeneity without F. M. Salzano which suggest8.6% if central reducing diversity of red cell antigens. Spain (Mourant et aI. 1976) were represen- However, the Brasilian tribes have been tative of the Caucasianintrusion. only minimally exposed to infectious Individual diversity in histocompatibility diseases introduced in post-Columbian antigens is restricted not only by the times, except malaria (Black et alo 1974). narrow range of alleles represented, but Furthermore, it now seems that other poly- also by the fact that a few haplotypes morphisms are reduced, there being charac- predominate. Four of these account for teristically only O of the ABO, reduced Rh more than half the sequencesfound in the and Gm diversity and no variation in Brasilian Indians. The variety of haplotypes Albumin, Hemoglobin or many enzymes found in the Kayapo and Tiriyo was much (Salzano 1972). The absence of whole greater than in the less bellicose Parakanã cross-reactive groups of HLA antigens and Waiãpi. We believe that this mar be would require that the selective pressure due to the practice of incorporating cap- operated across these groups. It seems tives into the more warlike ttibes. The more likely that the migrants to the conti- differences between me two Parakanã nent carried with them a limited spectrum villages and between the Kayapo villages of often related types. In as much as are pronounced. These differences are limited genetic diversity seems to have presumably due to genetic drift and serve preceded the onslaught of newly intro- to emphasize the power of this phenomenon duced infectious disease, it seems more in cultures of this kind. plausible that, if there is a relationship, it is The limited diversity of haplotypes to the cause rather than to the effect of would lead, according to the Hardy- diseasedepredation. Weinberg equilibrium, to the expectation that nearly 18% would be homozygous. Acknowledgment The number found, 12%, was unusually We are grateful to Dr. Amauri Azevedo and large but substantially (P = 0.02) lower many other members of the Fundação than expected. This discrepancy has been Nacional do Indio for transportation, trans-

374 HLA IN S. AMERINDIANS 375 lation and hospitality in Brasil; to Dr. J. M. (1974) The HL-A polymorphism in Mayan Borgono and the Servicio Nacional de Indians of San luan La Laguna, Guatamala. Histocompatibility Testing 1972, pp. 367- Salud for similar help in Chile; to Dr. 375. Munksgaard,Copenhagen. Francisco Pinheiro of Fundação Serviços 4:ann, H. M., Kidd, K. K., Lisker, R., Radvany, R. de Saude Publica for guidance and labora- & Payne, R. (1973) Genetic structure of the tory facilities; and to Dr. Zulay Layrisse of , HL-A system in a Nahua Indian population in Instituto Venezolano de Investigaciones Mexico. TissueAntigens 3, 364-372. Corley, R. B., Spees, E. K., Cabera, M. G., Científicas for evaluative and technical Swanson, I. L. & Amos, D. B. (1974) Histo- advice. compatibility Testing 1972, pp. 351-357. Munksgaard,Copenhagen. References Dossetor, I. B., Howson, W. T., Schlaut, I., Amos, D. B. & Ward, F. E. (1975) lmmuno- McConnachie, P. R., Alton, I. D. M., Lock- genetics of the HLA system. Pbysiol. Rev. 55, wood, B. & Olson, L. (1974) Study of the 206-246. HL-A system in tWo Canadian Eskimo Popu- Bashir, H. V., MacQueen, I. M., Amos, D. B., lations. Histocompatibility Testing 1972, Guinan, I. I., lohnston, I. M., Brotherton, pp. 325-332. Munksgaard,Copenhagen. I. v. M., Boettcher, B. & Ashton, G. C. Etcheverry, R., Guzman, C., Hille, A., Nagle, R., (1973) A study of the HL-A system in an Covarrubias, E., Regonesi, C., Muranda, M., Australian aboriginal population. Histo- Duran, N. & Montenegro, A. (1967) Investi- compatibility Testing 1972, pp. 311-315. gacion de grupos sanguíneosy otros caracteres geneticos sanguíneosen indígenas de Chile. I Munksgaard, Copenhagen. Black, F. L., Hierholzer, w. I., Pinheiro, F. P., En Atacamenos y Mapuches. Rev. Med. Chile Evans, A. 5., Woodall, I. P., Opton, E. M., 95, 599-604. Emmons, I. E., West, B. 5., Edsall, G., Downs, Faron, L. C. (1968) The Mapuche Indians 01 W. G. & Wallace, G. D. (1974) Evidence for Chile. Holt, Rinehart & Winston, New York. persistence of infectious agents in isolated Frikel, P. (1961) Fases culturais e aculturação human populations. Amer. J. Epidemiol. intertribal no tumucumaque. BoI. Mus. E. 100, 230-250. Coeldi, Belém 16NS, 1-90. Black, F. L., Pinheiro, F. P., Hierholzer, w. I. & Frikel, P. (1970) Os Kaxuyana. Notos etno- Lee, R. V. (1977) Epidemiology of infectious historicas. Pub. avulsas do Mus. E. Coeldi, disease: the example of measles. ln Healtb 14, 1-82. and Disease in Primitive Societies,pp. 115- Grofton, I. P., Chalmers, A., Price, G. E. & 130. Ciba 5ymposium 49, new series.Elsevier; Reeve,C. E. (1975) HLA 27 and ankylosing Amsterdam. spondylitis in B.C. Indians. J. Rheumatol. 2, Black, F. L., Pinheiro, F.P., Oliva, O., Hierholzer, 314-318. W. I., Lee, R. V., Briller, I. E. & Richards, lohnson, A. H., Noreen, H., Spees,E. K., Viii a- V. A. (1978) Birth and survival patterns in lobos, H., Serrano, H., Amos, D. B. & Yunis, numerically unstable proto-agricultural E. I. (1978) The distribution of HLA antigens societies in the Brasilian Amazon. Med. in the Motilones Indians of Venezuela. Tissue Antbrop. 2, 95-127. Antigens 12, 163-169. Black, F. L., 5a1zano,F. M., Layrisse, Z., Franco, Kissmeyer-Nielsen, F., Kjerbye, K. E., Lamm, M. H. L. P., Harris, N. 5. & Weimer, T. A. L. V., lórgensen, I., Bruun Petersen, G. & (1980) Restriction and persistence of poly- Gurtler, H. (1974) Study of the HL-A system morphisms of HLA and other blood genetic in Eskimos. Histocompatibility Testing 1912, traits in the Parakanãlndians of Brasil. Amer. pp. 317-324. Munksgaard,Copenhagen. J. Pbys. Antbropol. 52,119-132. Layrisse, Z., Layrisse, M., Heinen, H. D. & Bodmer, W. F. (1975) Evolution of HL-A and Wilbert, I. (1976) The histocompatibility other major histocompatibility systems. system in the Warao Indians of Venezuela. Genetics 79 suppl., 293-304. Science 194, 1135-1138. Cann, H. M., Bodmer, I. G. & Bodmer, W. F. Magalhães Santos, A. C. (1976) Os Parakanan. 376 BLACK ET AL.

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