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Interciencia ISSN: 0378-1844 [email protected] Asociación Interciencia Venezuela

Morales, Jorge Omar; Dipierri, José Edgardo; Alfaro, Emma; Bejarano, Ignacio Felipe Distribution of the abo system in the : miscegenation and genetic diversity Interciencia, vol. 25, núm. 9, diciembre, 2000, pp. 432-435 Asociación Interciencia Caracas, Venezuela

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DISTRIBUTION OF THE ABO SYSTEM IN THE ARGENTINE NORTHWEST: MISCEGENATION AND GENETIC DIVERSITY

Jorge Omar Morales, José Edgardo Dipierri, Emma Alfaro and Ignacio Felipe Bejarano

SUMMARY

Based on the ABO system phenotypes of 95,562 individuals and Black) was estimated. Although no significant interpopula- from the six that constitute the Argentine northwest tion differences of allele frequencies were detected, their geo- (Jujuy, , Tucuman, Catamarca, La Rioja, and Santiago del graphical distribution agreed with other sources of biological Estero), allele frequency, heterozygosis, and genetic admixture, information and with ethnohistorical and census data on this assuming three parental populations (American Indian, Spanish, region’s settlement.

Introduction groups, this polymorphism is (Lorandi and Otonello, 1992). the ANW population. They systematically and widely re- The Argentine northwest were: a) The pre-colonial Erythrocyte polymorphisms corded and documented, mak- (ANW) did not escape this stage (indigenous); b) The co- have exhaustingly been used ing it a traditional marker of process. It became a region lonial stage (Spanish and to characterize - both biologi- choice that supplies mass in- thanks to the singularity de- Black); and c) The national cally and ethnographically - formation on intra- and inter- rived from its physical, eth- State (other ethnic groups) different populations, and to population genetic variability. nic, historical and religious (Dipierri et al., 1996). reconstruct the history and Following the discovery of structure (Ruiz, 1993). Start- The aim of this study was geography of human genes America, new biological and ing from the time of arrival to determine the geographic (Comas, 1974; Mourant et al., cultural realities began to take of allochthonous extraconti- distribution of the ABO sys- 1976; Cavalli-Sforza et al., shape, with and Cre- nental populations to the re- tem in the ANW in order to 1994). Due to the ease for ole societies resulting from gion, three stages were deter- genetically characterize its ur- testing for the ABO blood mixes and new migrations mined in the constitution of ban mestizo populations, and

KEY WORDS / ABO / Miscegenation / Genetic Diversity / Argentine Northwest /

Jorge Omar Morales. Anthropol- Facultad de Humanidades y 4600 . cias Sociales. Universidad Na- ogist. Instituto de Biología de la Ciencias Sociales. Universidad . cional de Jujuy. Altura. Universidad Nacional de Nacional de Jujuy. Address: e-mail:[email protected] Ignacio Felipe Bejarano. Anthropol- Jujuy. Instituto de Biología de la Altu- Emma Alfaro. Geneticist. Instituto ogist. Facultad de Humanidades José Edgardo Dipierri. MD. Insti- ra, Universidad Nacional de de Biología de la Altura y Fa- y Ciencias Sociales. Universidad tuto de Biología de la Altura y Jujuy, Avda. 1665, cultad de Humanidades y Cien- Nacional de Jujuy.

432 0378-1844/00/09/432-04 $ 3.00/0 DEC 2000, VOL. 25 Nº 9 RESUMEN

Con base en los fenotipos del sistema ABO de 95.562 indi- Si bien no se detectaron diferencias significativas en las fre- viduos de las seis provincias del noroeste argentino (Jujuy,Salta, cuencias de alelos entre poblaciones, su distribución geográfica Tucumán, Catamarca, La Rioja y ) se estimó fue concordante con otras fuentes de información biológica y la frecuencia alélica, la heterocigosis y la miscegenación, asu- con datos etnohistóricos y censales de los asentamientos huma- miendo tres poblaciones parentales (india, española y negra). nos en esta región.

RESUMO

Com base nos fenotipos do sistema ABO de 95.562 indiví- Se bem que não detectaram diferencias significativas nas fre- duos das seis províncias do nordeste argentino (Jujuy,Salta, qüências de alelos entre populações, sua distribuição geográfi- Tucumán, Catamarca, La Rioja e Santiago do Estero) se esti- ca foi concordante com outras fontes de informação biológica e mou a freqüência alélica, a heterocigosis e a miscegenação, com dados de etnohistóricos e censos dos assentamentos huma- assumindo três populações parentais (índia, espanhola e negra). nos nesta região.

to analyze the biological af- through time as it is the terri- finities among them and their tory that best resisted the proc- degree of genetic admixture. ess of denaturalization caused by successive ethnic disrup- Populations and Methods tions that took place during the different stages of settle- The ANW extends between ment of the region. 22º 46' and 30º11' southern Sources for the data used in latitude, and between 61º43' this study were: a) blood do- and 69º 05' western longitude. nors; b) records of driving li- It consists of six provinces cense applicants; c) incorpora- (Figure 1) with a total popu- tions to military post stations lation of 3,677,538 inhabit- in the ANW. Determinations ants. From the biogeographi- for each included cal point of view, it is a sec- genotype and alleles frequen- tion of the Neo-Tropical Re- cies of the system (maximum gion, with a wide diversity of likelihood method, MAXLIK atmospheres (mountains, high software) heterozygosis (Nei, altitude deserts, and forests) 1973), and genetic admixture (Cabrera and Willink, 1973). (Chakraborty, 1985), assum- Earliest cultures inhabiting ing three parental populations: the current ANW were limited a) Spanish (Junyent et al., to some ethnical groups that 1992); b) South-American In- linguistically used a mosaic of dians (Salzano and Callegari- languages and dialects (Gon- Jacques, 1988); and c) Black zález and Pérez, 1972). From (Castro de Guerra, 1993). the cultural point of view, Nei’s method (1973) was they belonged to the North- used to determine genetic dis- west Cultural Area, Sub-Area tances and the corresponding of the South Andean Area tree was developed from and macro Andean Area there. As no statistically sig- (Hernández, 1992). It consti- nificant differences were ob- tutes the most important cul- served among the alleles fre- Figure 1. Mapa tural area in the Argentine ter- quencies from the various ritory with the most relevant sources, estimates were made consequences in the develop- by grouping them. (0.81±0.001), followed by A Estero showed the highest AB ment of its population. This (0.14 ± 0.001) and B (0.05 ± penotype percentages. characteristic is partially deter- Results 0.001) (Table I). Allele fre- Average heterozygosis (AH) mined by: a) The numerical quencies did not present sta- among populations was of relevance of its native popula- The most frequent pheno- tistically significant interpro- 0.321, while total heterozygosis tion, for it registered the type we found over a total of vincial differences. The prov- was of 0.326, and interpopula- highest population density in 95,562 individuals was 0 inces with highest percentages tion genetic diversity was of pre-Spanish times (Hernández, (66%), followed by A (24%), of 0 phenotype, however, 0.015. The highest heterozygo- 1992); b) The permanence of B (8%), and AB (2%). The were Jujuy and Salta, while sis was found in Tucumán, and this autochthonous population most frequent allele was 0 Tucumán and Santiago del the lowest, in Jujuy (Table II).

DEC 2000, VOL. 25 Nº 9 433 TABLE I PHENOTYPES AND ALLELE FREQUENCIES

PROVINCES JUJUY SALTA TUCUMAN STGO. CATAMARCA LA RIOJA TOTAL DEL ESTERO

Phenotypes N % N % N % N % N % N % N % A 1199 14.66 4789 16.92 3479 30.72 8896 28.01 2907 30.49 1839 28.43 23109 24.18 B 476 5.82 1542 5.45 1147 10.13 3055 9.62 863 9.05 474 7.33 7557 7.91 AB 48 0.59 214 0.76 255 2.25 1000 3.15 185 1.94 112 1.73 1814 1.90 O 6453 78.93 21758 76.87 6442 56.90 18805 59.22 5580 58.52 4044 62.51 63082 66.01

Allele Frequencies A 0.08 ± 0.002 0.09 ± 0.001 0.18 ± 0.003 0.17 ± 0.002 0.18 ± 0.008 0.16 ± 0.003 0.14 ± 0.001 B 0.03 ± 0.001 0.03 ± 0.001 0.06 ± 0.002 0.07 ± 0.002 0.05 ± 0.003 0.05 ± 0.002 0.05 ± 0.001 O 0.89 ± 0.002 0.88 ± 0.001 0.76 ± 0.003 0.76 ± 0.002 0.77 ± 0.007 0.79 ± 0.004 0.81 ± 0.001

A high proportion (above would result from intrapopu- and persisted with the arrival sult, these populations were 60%) of autochthonous mix- lation differences. of the Spanish conqueror, severely ravaged and denatu- ture was observed in the prov- The highest frequency of 0 whose influence altered popu- ralized (Lorandi and Otonello, inces of Jujuy and Salta, while allele was found in the prov- lation dynamics and the eth- 1992; Hernández, 1992). The Spanish contribution (above inces of Jujuy and Salta nic relationship through dif- opposite took place in the 40%) was found in Catamarca, (Table I). Coincidentally, ferent mechanisms of mass current provinces of Jujuy La Rioja and Tucumán. The these provinces also showed denaturalization of natives and Salta, where rebellion Black component (over 20%) lower average heterozygosis, (mitimaes, yanaconazgos or was shorter and communities was found in Santiago del higher biological affinity with Indian binding to personal adopted resistance strategies Estero, Tucumán, and the American Indian parental service, wars, and other forms and reached agreements with Catamarca (Table II). The population (Figure 2), and a of migration and accultura- the conqueror that allowed populations of Jujuy and Salta higher percentage of native tion) (Bazán, 1986; Lorandi them to mitigate the effects of constitute a subgroup with the admixture (Table II). A pre- and Otonello, 1992; Hernán- denaturalization (Bazán, 1986; American Indian parental liminary explanation of these dez, 1992). This denaturaliza- Lorandi and Otonello, 1992). population, while the rest of results could relate to the dif- tion process was more intense The process continued with the populations were grouped ferential ethnic disruption of in ethnic groups (, the flow of Black people. Ac- together with the Black popu- pre-Columbian native commu- Lules-Vilelas, Quilmes, Tono- cording to Goldberg and lation, leaving the Spanish nities of the ANW and the cotes) that populated the cur- Mallo (1993), between 1740 population in a separate sub- characteristics of the migra- rent provinces of Tucumán, and 1810, 45,000 African in- group (Figure 2). tion process in this region La Rioja, Catamarca, and dividuals entered through the during the XIX and XX cen- Santiago del Estero, which port of and dis- Discussion turies. from 1561 to 1655 partici- persed over the colonial Ar- Disruption started with the pated in the rebellions against gentine territory. This genetic Although this study only arrival of the Inca conqueror Spanish domination. As a re- flow determined that during used a polymorphic system, it provided information on ge- netic variability of an impor- TABLE II tant percentage (2.7%) of the PERCENTAGES OF GENETIC ADMIXTURE AND AVERAGE HETEROZYGOUSITY ANW population. Available literature includes few local and national studies on the PROVINCES PARENTS AVERAGE ABO system (Scaro, 1958; POPULATIONS HETEROZYGOUSITY Quiroga Micheo et al., 1988) AMERINDIAN SPANISH BLACK and none of them approaches the estimation of allele fre- JUJUY 0.71 0.15 0.14 0.204 quencies. SALTA 0.67 0.23 0.10 0.225 Broadly results indicate that TUCUMÁN 0.28 0.46 0.26 0.393 there is scarce interpopulation heterogeneity and low abso- SGO. DEL ESTERO 0.30 0.39 0.31 0.380 lute and relative genetic di- LA RIOJA 0.40 0.49 0.11 0.383 versity in the ANW (Table CATAMARCA 0.32 0.48 0.20 0.347 II), as 99.9% of total genetic TOTAL 0.45 0.34 0.21 0.322 variation found in this region

434 DEC 2000, VOL. 25 Nº 9 uted to this study, the direc- Dipierri JE, Alfaro E, Peña JA, JUJUY tors of provincial Traffic Con- Jacques C, Dugoujon JM trol Offices, and the Chief of (1999b) GM, KM immunoglo- SALTA bulin allotypes and other se- Recruitment and Mobilization, rum genetic markers (HP, GC, AMERINDIANS Jujuy Division, of the Argen- PI and TF) among South tine army, without whose col- American populations living TUCUMAN laboration in the provision of at different altitudes (, Argentina) Admix- data this study would not ture estimates. Hum. Biol. CATAMARCA have been carried out. 72(2):305-319. SANTIAGO DEL ESTERO Goldberg M, Mallo S (1993) La población africana en Buenos LA RIOJA Aires y su campaña. Formas REFERENCES de vida y de subsisten- BLACK cia.1750-1850. Facultad de Filosofía y Letras. Universidad Bazán AR (1986) La Historia del de Buenos Aires (manuscrito). SPANISH Noroeste Argentino. Plus Ul- tra. Buenos Aires. González A, Pérez J (1972) Argen- tina Indígena. 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These showed high and other animals popula- tions. Indian Anthropological The distribution of the human of genetic admixture (Table frequency of typically Ameri- Association. blood groups and other ge- II) and it would explain the can Indian markers, a signifi- netic polymorphisms. Oxford greater affinity of the popula- cant component of native Comas J (1974) Antropología de University Press, London. los pueblos iberoamericanos. Nei M (1973) Analysis of gene tions in Tucumán, La Rioja, mixture in all the provinces, Biblioteca Universitaria Labor. Santiago del Estero, and mainly those of Jujuy and diversity in subdivided popu- Barcelona. lations. Proc. Natl. Acad. Sci. Catamarca with their Black Salta, a high percentage of 70: 3321-3323. parental population (Figure Black mixture and scarce ge- Dipierri JE, Alfaro E, Bejarano I 2). netic interpopulation heteroge- (1996) El Patrimonio Bio- Quiroga Micheo E, Villaseca A, lógico de la Provincia de Ju- Bonder M, Quiroga Vergara E During the stage of confor- neity (Dipierri et al., 1997, juy. Revista XUXUY. Ciencia (1988) Frecuencia de los mation of the national State 1999b). & Tecnología. 1(2): 13-21. grupos sanguíneos y análisis (by the end of the XIX cen- ABO system characteristics SECTER - UNJu. de la progresiva disminución tury and turn of the XX cen- found in the ANW point to a del factor RH negativo en la Dipierri JE, Alfaro E, Vullo C, población Argentina. Medicina tury), data provided by the variation that agrees with the Gutierrez N (1997) El poli- (Buenos Aires) 48:355-366. 1914 Census (Tercer Censo behavior of other genetic morfismo HLA en el noroeste Nacional, 1914) showed that polymorphisms indicative of argentino. III Jornadas Nacio- Ruiz M (1993) Prehistoria y nales de Antropología Bioló- arqueología regional. En: the provinces of Salta, Tucu- the small degree of genetic gica. Rosario. Argentina. Campi D (Ed) Jujuy en la mán, Santiago del Estero, and differentiation in the popula- historia. Avances de Inves- La Rioja received a larger tions of this region (Dipierri et Dipierri JE, Alfaro E, Martinez tigación. Unidad de Historia number of European individu- al., 1998; 1999a). They also Marignac V, Bailliet G, Bravi Regional. FHyCS.UNJU CM, Cejas C, Bianchi NO als. It doubles or triples the agree with a miscegenation (1998) Paternal directional Salzano F, Callegari-Jacques S number of those who settled process related to the popula- mating in two amerindian sub- (1988) South American Indi- down in Jujuy, depending on tion development stages populations located at differ- ans. A case Study in Evolu- ent altitude in the northwest tion. Clarendon Press. Oxford, the province. The opposite shown by historical and de- USA. happened with individuals of mographic sources. of Argentina. Hum. Biol. 79 (6): 1001-1010. Scaro J (1958) Investigación del American origin, mainly rep- Factor Diego en aborígenes de resented by Bolivians, which ACKNOWLEDGEMENTS Dipierri JE, Alfaro E, Bejarano I la Quebrada de . constitute 74% of migrants to (1999a): Surnames, AB0 sys- Rev. Soc. Arg. Biol. 34:71-75. tem and miscegenation in Jujuy, 34% to Salta, while it The authors thank the au- highlands population of Prov- Tercer Censo Nacional (1914). is non-existent or under 1% thorities of the various hospi- ince of Jujuy (Northwest Ar- Talleres Gráficos LJ Rosso, in other provinces. Once tals that generously contrib- gentine). HOMO 50(1): 14-20. Buenos Aires.

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