HLA Antigens in South American Lndians
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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 (Wayampi, 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 Iarge 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 South America 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} Can .b 17.Mapuche18.Ticuna (approx. 38o S 71 oW) 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.