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Genetic variation in the Bribri and Cabecar Amerindians from Talamanca, Costa Rica

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(1)Rev. Bio!. TI'Op 39 (2): 249-253,1991 .•. Genetic variation in the Bribri and Cabecar Amerindians from Talamanca, Costa Rica Jorge Azofeifa y Ramiro Bammtes. Instituto de Investigaciones en Salud (lNISA) y Escuela de Biología, Universidad de Costa Rica, Costa Rica , América Central. (Rec. 28-IX-1990. Acep.5-IV-1991). in Ihe Bribri and Cabecar Amerindian populations of Talamanca in Soutbeastem Costa Rica. Sorne differences were found in fue distributiOll (presence or absence) oí Ihe variants (some oí fuero private) oí Ihe GOT. PEPA, RH. J'F and TPI loo when Ihe results reported bere were compa­ red wilh Ihose oí Ihe Bribris and Cabecars that inhabit in Ihe Pacific side of Ihe country. Admixture wilh non-Indians was detected at Ihe ABO, ADA, and G6PD loci. The proportion of poIymorplllc loo (P) found were 17.1 in the Bribris and 19.4 in Ihe Cabecars; Iheir average beterozygosities (H) estimates were 0.039 and 0.049 respectively. Fst values (0.019 in Ihe Bribris and 0.056 in the Cabecars) seero to indicate Ihat random events have pIayed important ro­ les in the divergence of Ihe Cabecars. Low effective population sizes oí the emignmt groups to Ihe Pacific side and probable gene flow in fue case of Ihe Bribris, could be Ihe causes oí Ihese differences.. Abstrad: A screening of more Ihan 40 loo was penormed. Key words: Amerindians, genetic variation, polymOIphic loci, beterozygosis. geographic variation.. The importance of studying Amerindían groups lo understand sorne aspects of human evolution was pointed out by Neel and Salzano (1964); after that, a great de al of work was per­ formed particularly in South America (Salzano & Callegari-Jacques 1988). In Lower Central America, systematic work began by 1919 with the study of the Costa Rican and Panamanian groups (Spielman et al. 1979, Barrantes et al.. 1982, 1990).. The Bribris and Cabecars represent together more tlmn fúty percent of the total Costa Rican Amerindian population that was estimated to be around 13000 by 1982 (Bozzoli 1986). These groups are found in the south of the country, on both sides of the Cordillera de Talamanca (Talamanca Mountain Range): ne­ arly sixty percent in the Atlantic and !he remai­ ning in fue Pacific (Bozzoli 1986), whose for­ mer settlers were irnmigrants from the southe­ astero (Atlantic) part of the country (Stone 1962). Genetic studies include a previous exa­ mination (Azofeifa 1987) of part of fue same sample here reported, and a phylogenetic analysis of fue Chibcha-speaking groups of. Costa Rica and Panama (Barrantes et al. 1990). Here we present new information on nearly 40 genetic loci that show that their genetic structu­ res are similar to those of other Amerindian groups of Lower Central America (Barrantes el al. 1990) and South America (Neel 1978, Salzano & CaUegari-Jaques 1988), although the gene frequencies distributions as well as fue presence of private variants at certain loci are distinctive within each group.. MATERIAL AND METHODS Samples were collected during three trips 10. fue field. The flfSt two were lO the Talamanca Indian Reservation that is located between the 9" 27' and 9" 40' N and between 82° 50' and 83 , .7' W, at the localities of Amubri, on May,. 1984 and Mojoncito, on February, 1985. The Bribris and Cabecars in fuese pIaces are here referred as Talamancans. Too third trip was lO a place near the locality of Moravia in the Chirripo Indian Reservation, between !he 9° 43' and 9° 54' N and between 83° 32' and 83° 36' W on February, 1986..

(2) RBVISTA DB BIOLOGlA 'IROPICAL. �lood was obtamed, transported and proces­. sed as described elsewbere (Barrante s et a l.. 1982). The nwn� of samples analysed, accor­ diog 10 lOO �lared etbnie group of lOO donors , is shown in Table l. Differences in these figu­ res are due lO availability of reagents al the mo­ ment of tbe sampling and lO damage of storing containers. Tbe ABO, Duffy, MN, RH and Ss blood group typings were performed by tbe saline tu­ be test and following lOO antisera manufactu­ rers' indieations. T be electtopboretie anaIyses were done using botb polyaerylamide and starcb gels. In polyacrylamide tbe following 10ei were screened: a. enzylJ'l.es: adenosine .dea­ minase (ADA) glueose pbosphate isomerase (GPI). lactate debydrogenase (LDH), malate debydrogenase (MDH), 6-pbospboglueonate dehydrogenase (PGD) and triose pbosphate iso­ merase (TPI); b .serum proteins: albumin (ALB), eeruloplasmin (CP), haplOglobin (HP) and transferrin (TF); e.bemoglobins: A (HBA) and A2 (HBA2) following tbe same metbods employed by os in otber works (Bammtes el al 1982, 1990, Bammtes, Azofeifa & Mata 1985). In starcb geIs tbe following enzymes were stu­ died: acid phosphatase (ACP1), adenylate kina­ se (AK1), carbonie anhydrases 1 and 2 (CAl, CA2), NADH diapborase (DIAl), esterases A, B, C and D (ESA, ESB, ESC, ESD), glutamate­ oxaloaeetate transaminase (GOT), isocitrate debydrogenase (ICD), lactate debydrogenase (LDH), malate debydrogenase (MDH), nucleo­ side pbospliorilase (NP), peptidases A, B, C and D (pEPA, PEPB, PEPC, PEPD), phospho­ glueomutases 1 and 2 (PGM 1, PGM2) and su­ petOxide dismutase (SOD) using tbe techniques described in Bammtes et al., ( 1982 , 1990) and Harris and Hopkinson (1976). For glucose-6phosphate dehydrogenase the metbodology used w as that detailed by Motulsky and Yoshida ( 1969) . The allele frequeney calculations were done by tbe gene counting metbod. The average he­ terozygosities (gene diversities) (Nei 1987) and proportions of polymorpbic loci were estimated for tbe 35. (Bribris) and 36 (Cabecars) electrop:' hoteticallystudied loci. The criterion we osed for a polymOtphic locos was that tbe frequency of its commonest allele does not exceed 0.99. To estiniate tbé effect of population subdivi­ sioR, Fst values, based on tbe observed and ex­ pected beterozygosities (Nei 1987), were calcu-. Jated using 6 (Bribris) and 8 (Cabecars) poly­ morpbic loci (tbe phenotype data on wbich the­ se estimators were based could be sent on re­ quest). The variation detected witb non-Indian alleles was nOl included in tbe estimations.. RESULTS. The allelic frequencies of lOO genetic sys­ tems at wbich variation was observed are shown in Table 1. Most of tbe loci analysed by electtophoresis were foundlO be monomor­ pbic, what .lpeans in terms, of P (proportio n of polymorpbic loci) 17.1% in tbe Bribris and 19.4% in tbe Cabecars. The average heterozy­ gosities (H) obtained were 0,039 and 0.049 res­ pectively. Racial admixture witb non-Indians was evident al tbree loci: al tbe ABO blood group system, at tbe ADA enzyme locos and al tbe G6PD enzyme locus. In general, three groupings can be made witb tbe variant loci: a. tbose with alleles normally foundin almost every studied population (Harris & Hopkinson 1976) and tbat present, basically, tbe same allele frequencies in tbe groups compared bere; tbey are ACPl and PGM 1. b. Five loci, ESD, FY, JIP, MN and Ss show variation witb segregating alleles common 10 nearly all the hwnan populations (Harris & Hopkinson 1976, Mestriner, Simoes & Salzano 1980, Bowman & Kurowsky 1982, Mourant 1983) but witbout a clear pattem among tbe groups compared in Table 1. c. A third group of loci shows a rather interesting variation tbal in­ cludes. besides lOO normal alleles, private va­ riants restricted lO Amerindian populations. or 10 otbers related 10 them. The notability oí tbese loci comes from tbe presence or absence of a determined allele in the Bribri and Cabecar·po­ pulations from tbe Pacific side of tbe Talamanca Mountain Range and tbe Bribri and Cabecar onesfrom its Atlantic side. They are GOT, PE­ PA (that sbows. besides the normal. a low-acti­ vity variaIit allele, PEPA * 2KUNA up lO now detected onIy in tbe Chibcha-spealdng groups of Lower Central America; Bammteset al. 1990), RH, 1F (tbat includes two variant alleles, D­ China on TF*DCHI, an allele disttinctive of Mongoloids and Amerindians; Neel 1978; as well as D-Guaymi (TF*DGUA), that was f11"st detected in a sample of Panamanian Guaymis; in Tanis, Neel & Torres � Arauz 1977) and 1PI (wbere tbe private 1PI*3BRI allele was ob-.

(3) 251. A ZOFEIFA &. BARRANTES: Genetíe variauro in Amerindias fmm Costa Rica. TABLE 1 Allelicfrequenciesfor the systems thal showed varia/ion in the Bribri and Cabecar from lhe Atlantic o/Costa Rica. and a comparison wilh Ihose olother Costa Rican and Panamanian Amerindians*. Pacific. A tlanuc. Gy(TT O). Locus. A llele. Br(123). Ca(99). Br(99)@. Ca(60)@. Gt(80)@. ABO. Al A2 O A B BGUA 1 2 1 2 a b 1 2 3 A B. 0.004 0.004 0.992 0.131 0.869. 1.000 0.077 0.923. 1.000 0.131 0.869. 1.000 0.075 0;925. 1.000 0.084 0.160. 1.000 0.103 0.897 (+C). 0.990 0.010 0.912 0.088 0.931 (58) 0.069 0.991 (57) 0.009. 1.000. 1.000. 1.000. 1.000. 0.939 0.061 0.641 0.359 0.949 0.036 0.015. 1.000. 0.717 0.283 0.526 0.474 1.000. 0.690 0.310 0.373 0.627 0.952 0.032 0.016. 0.958 0.042 0.478 0.522 0.988 0.012. 1.000 0.383 0.671 0.919 0.081 1.000. 1.000 0.525 0.475 0.975 0.025. 0.908 0.092 0.200 0.467 0.333. 0.460 0.540 0.853. 0.450 0.550 0.966. 1.000 0.661 0.339 0.786 0.214 0.794 0.206 0.921 0.079 0.032 0.413 0.547 0.008 0.500 0.500 0.794 0.206. 1.000 0.545 0.455 0.686 0.314 1.000. 0.924 0.076 0.030 0.546 0.424. 1.000 0.410 0.590 0.506 0.494 0.699 0.301 0.964 0.036 0.050 0.519 0.418 0.013 0.234 0.766 1.000. 0.147 1.000. 0.034 1.000. 0.976 0.024. 1.000. 1.000. ACPI. A DA ESD FY GOT. G6PD. 1.. HP MN PEPA PGMl RH. 2 M N N 2KUNA 1 2 O 1 2. Notes:. @. ND 1'>.'1) ND 0.009 0.991 0.516 0.484 ND ND 1.000. DGUA 1 3BRI. 0.952 0.048. C. TPI. ND. DCill. S. TF. 0.893 0.107 ND. 0.984 0.016 0.079 0.540 0.357 0.024 ND ND 0.837 0.163. Z. Ss. 1.000. Te(63)@. (MB) 0.002. 1.000 0.321 0.679 0.741 (58) 0.259 0.760 0.240 0.897 0.103 0.052 0.448 0,490 0.010 0.810 (58) 0.190 0.747 0.016 0.237 0.974 0.026. 0.808 0.192 0.975 0.025. 1.000. 0.998 0.057 0.811 0.132 1.000. 0.947 0.053 0.023 0.845 0.132 0.380 0.620. 0.945 0.055. Sample sizes indicated in brackets . Br=Bribri, Ca=Cabecar, Gt=Guatuso, Te=Terihe, Gy=Guayrni, ND=no data.. Does not include allthe vanant loo found in the groups fmm outside Talamanca III1d Chirripo. Frequencies calculated fmm lhe gene counts reporte<! by Barrantes el al. (1990). The frequencies of lhe Guayrnis from Barrantes el al., (1982).. are.

(4) 252. REVISTA DE BIOLOGIA TROPICAL. served). This variant (Figure 1) that was frrst fmmd in the Cabecars of Talari (Barrantes e t al. 1990) is characterized by 1ts particular stai­ ning (activity) pattem rather than by an altcred mobility of any specific band, and seems ro be restricted, witb tbe exception of tbe Guatusos, ro the Talamancan groups.. Figure 1 . Electrophoretic pattems oí me TPI 1.1 (nonnal), lanes 1 ,2 and 4, and TPl 1 , 3 BR!, la.nes 3, 5 and 6 alter a PAGE (7% acrylamide , pH 8.5). Note !he stronger acti­ vity oí bands lA b and e in me nonnal and of bands e d , e and f in me heterozygQ(e wim me variant. •. •. Fst values were 0.019 in the Bribri and 0.056 in the Cabecar. DISCUSSION T he genetic structures of the Bribri and Cabecar populations of Talamanca are very si­ milar ro those of other Amerindian populations (Barrantes el al. 1982, 1990, Neel 1978) as can be seen from the high proportion of mono­ morphic loci, the low values of average hete­ rozygosities (H) as well as for the presence of at least o n e private p o lymorphism, the TPI*3BRI and the PEPA *2KUNA aUdes, in both groups considered here. As expected, dif­ ferences in gene frequencies were observed between the groups, particl.llarly if we take into accannt their estimated time of divergence, that ranges, according ID lexicostatistical and gene­ tic analyses, from 600 to 1500 years (Barrantes et al. 1990, Bozzoli, 1979, Constenla 1985, Vargas 1986). An early observation (Azofeifa 1987) detec­ ted differences between the Bribris o f Talamanca and the Bribris o f Salitre, and bet­ ween the Cabecars of Talamanca and the. Cabecars of Ujarras. They were evi<knt when we frrst comoared most of our data with those of Matson Md his groop (Matson & Swanson 1965, Matson el al. 1965) which served as a source for the Pacific side populations. These desimilarities are now confmned wnen we use OUT group's data from Cabagra and Ujarras (Barrant es el al. 1990). In this latter case more . loci were compared. which were sampled and anaIysed nsing the same methodologies. The differences are specia1ly clear at the PEPA, TF, TPI and RH loci. Similar differences were found when the dermatoglyphic pattems of the­ se populations were compared (Quesada, M . & R. Barrantes, in preparation). The Fst vaIues of both groups suggest tbat different evolutionary determinants may have affected them. bringing the Cabecars to diverge more between them than the Bribris. The ran­ dom effect that results from subdivision, seems ID have played a main role in the Cabecars ; low effective population sizes of the emigrant gronps could explain this. Another important factor lhat could account fol' the dissimilarities observed. particularly in the Bribris, is admix­ ture with the Cabecars and other indian groups (Borneas, Tecibes). Thls has not been documen­ ted in the pasto but detected by demographic in­ formation (Azofeifa 1987, Barrantes & Azofeifa 1983). Admixture with non-Indians was also inferred due to !he resullts of certain lod. Despite the exclusion of bearers ol non­ Indian markers. an absolute depuration of the sample can never be guaranteed. ACNOWLEDGEMENTS To M . Chaves of the CIHATA, UCR, who provided !he control for G6PD*A . To J . Lobo and A . Chinnock for the critica! review of the manuscript . To L Castro and J . Gamboa for their help in the laboratory. This work was sup­ ported by project No. 111-79-005 of the University of Costa Rica. .. RESUMEN Se analizaron más de 40 locÁ de las poblacio­ nes amerindias bribri y cabécar de Talamanca, del sureste de Costa Rica . Se encontraron algu­ nas diferencias en la distribución (presencia o ausencia) de las variantes, algunas de elms pri­ vadas, de los loci de la GOT, la PEPA, el RH, la.

(5) 253. AZOFEIFA &. BARRANTES: Genetie vanatioo in Amerindias fmm Costa Rica. 1F Y la TPI. al compararse nuestros resultados. con los descritos para las. poblaciones bribris y cabécares de la región del Pacífico Sur del país. Se encontró mezcla con no indígenas por mter­ medio de los loci del ABO, de la ADA y de la G6PD. Los valores de la proporción de loci po­ limórficos (P) hallados fueron 17.1 en los bri­ · bris y 19.4 en los cabécares; las estimaciones de heterocigosis media (H) fueron 0.039 y 0.049 respectivamente. Los valores de Fst obte­ nidos (0.019 en los bribris y 0.056 en los cabé­ caces) sugieren que eventos aleatorios han sido más importantes en la divergencia de los cabé­ caces entre sí . Tamafios efectivos pequefios de los grupos emigrantes alIado del Pacífico y flu­ jo génico, en el caso de los bribris, pueden ser las causas de estas diferencias. REFERENCES. Const.enJa, A. 1 985. .. ClasificaciÓllleJÚcoes�stica de lu. lenguas de la Familia Chibcha. p. 155-1 97. In Departamento de Lingüística, Universidad de Costa Rica. E studios de Lingüística Chibcha , '101.4. Universidad de Costa Rica, San José. Harria, H. &. D.A. Hoplrinsoo . 1976 Handbook oí Enzyme Electrophoresis in Human Genetics. Norih Hollimd , Amst.erdam. •. Matsoo, G.A.. H.E Sutton, J. SWllDSon &; A.R. Robinsoo. 1965. Distributioo of haptogJ.obin, ttansferrin and he­ mogJ.obin types amoog Inmans of Middle America : In British Honduras,.Costa Rica and Panama. Am. J. Phys.. Anthrop. 23:123-13O. Matsoo. G.A. &. J. Swansoo. 1965 . Distributioo oí heredi­. tary blood antigens amoog Indians in Middle Amenca. vn. In Costa Rica. Am. J. Phys. An!hrop. 23:10"7-122. Mestrlner, M.A., A.L. Simoes &; EM. Salzano. 1980 .New studies 011 lhe esteraSe D polymorphism in South American Indians. Am. J. Phys. Anthrop. 52:95-1Ol.. Azofeifa , J. 1981. Estructura Genética y Demografía de la Población Amerindia de Talamanca ,Costa Rica M. Sc. Thesis . Universidad de Costa Rica .. 'Motulsky, A.C. &; A. Yoshida. 1969 . Methods fOI!he study of red cell glucose-6-phosphate dehydrogenase. p.5193. In lJ. Yunis (ed.). Biocherrucal Me!hods in Red eell Genetics. Academic Press , Loodoo.. Barrantes, R. &. 1. Azofeifa. 1983. Biodemografía de va­ rias poblaciones indígenas de Costa Rica . Vínculos 9:15-23.. MOUTant , A.E. 1983 . Blood Rciations. Blood Groops and Antbropology. Oxford, Thetford. .. Barrantes , R . ,1. Azofeifa &. L . Mata. 1985 . Grupos san­ guíneos ABO y Rh y proteínas séricas en una población amerindia , Matambú , Costa Rica .. Rev. Biol.Trop. 33:13-16. Barrantes, R, p.E. Smoose, J.V.Neel, H.W. Mohrenweiser, &. H. Gershowitz. 1982 . Migratioo and genetic infras­ tructure oC !he Central. American Guayrni and !heir af­ fmilies wiili other tribal groups. Am. J. Phys. An!hrop. 58:201-214. B arrantes, R., P.E. Smouse, H.W. Mohrenweiser, H.Gershowitz, J. Azofeifa, T.D. Arias &; J.V. Neel. 1990 .Microevolution in Lower Central America: G enetic characterization of ahe Chibcha-speaking groups of Costa Rica and Panama, and a concensus ta­ lI.ooomy based on genetic and linguistic affinities. Am. J. Hum. Geuet. 46:63-84.. Neci, J.V. 1978 . Rare variants, private polymorphisms and locos heterozygosity in Ametindian populatioos. Am. J. Hum. Gmet. 30:465-490. Neel, J.V. &. P.M. Salzano . 1964: A prospectus for genetic studies of the American Indian. Cold Spring Harbur Symp. Quant. Bio!. 24:85-98. Nei, M. 1981. Molecular Evoluliooary Genetics. Columbia University Press ,New York. Salzano, P.M. &. S.M. Callegari-Jacques. 1988 . Soutb American Indians. A Case Study in Evolucion. Oxford University Press ,New York. Spielman, RS., E.e. Migliazza, IV. Neel, H. Gershowitz &; R. Torres de Aiauz. 1979 The evolutionary rela­ tionships of two populations : A study oí !he Guaymi and Yanomama. Curro An!hropol. 20:317-388.. Bowman, B.H. &. A. Kurowsky 1982 Haptoglobin : tbe evolutiooary product of duplication , unequal crossing over and point muta1Íon. Adv. Hum. Gene!. 12:189261.. Stooe, D. 1962 . Las Tribus Talamanqueñas de Costa Rica . Lehman , San José "Costa Rica.. Bozzoli, M.E. 1919 . ElNacimiento y la Muerte entre los Bribris. Editorial Universidad de Costa Rica, San José.. Tarus, RJ., J.V. Neel &. R. Torres de Arauz. 1977 . Two more "private" polymorphlsms of Amerindian tribes : LDH B GUA-l and ACPl B GUA-! in !he Guayrru in Panama. Am. J. Hum. Genet. 29:419-430.. B ozzoli, M.E. 1986 El Indígena Costarricense y su Ambiente Natural: usos y adaptaciones. Editorial Porvenir, San José, Costa Rica.. Vargas, P. 1986 . Análisis Lexiooestadístioo de las Relaciones entre el Bribri y el Cabécar. Lic. Tesis.Universidad de Costa Rica..

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Figure

Figure 1 .  Electrophoretic pattems oí me TPI  1.1  (nonnal),  lanes 1 ,2 and 4, and  TPl 1 ,  3 BR!,  la.nes 3, 5 and 6  alter  a  PAGE  (7% acrylamide , pH 8.5)

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