SciELO - Scientific Electronic Library Online

vol.22 issue2Genetics, surnames, grandparents' nationalities, and ethnic admixture in Southern Brazil: Do the patterns of variation coincide?Diagnosis of the fragile X syndrome in males using methylation-specific PCR of the FMRI locus author indexsubject indexarticles search
Home Pagealphabetic serial listing  

Services on Demand


  • English (pdf)
  • English (epdf)
  • Article in xml format
  • Article references
  • How to cite this article
  • Curriculum ScienTI
  • Automatic translation
  • Send this article by e-mail


Related links


Genetics and Molecular Biology

Print version ISSN 1415-4757
On-line version ISSN 1678-4685

Genet. Mol. Biol. vol.22 n.2 São Paulo June 1999 

Genetical-Demographic Data From Two Amazonian Populations Composed of Descendants of African Slaves: Pacoval and Curiau

João Farias Guerreiro1, Ândrea Kely Campos Ribeiro-dos-Santos1, Eduardo José Melo dos Santos1, Antonio Carlos Rosário Vallinoto1, Izaura Maria Vieira Cayres-Vallinoto1, Gilberto Ferreira de Souza Aguiar2 and Sidney Emanuel Batista dos Santos1
1Laboratório de Genética Humana e Médica, Universidade Federal do Pará, 66075-970 Belém, Pará, Brasil. Send correspondence to J.F.G. Caixa Postal 8615. Fax: +55-91-249-0373. E-mail:
2MCT-PR/CNPq. Museu Paraense Emilio Goeldi, Departamento de Ciências Humanas, Belém, PA, Brasil.



The Amazon region of Brazil includes communities founded by escaped slaves, some of which still remain relatively isolated. We studied two such Afro-Brazilian communities (Pacoval and Curiau), in the rural area of Alenquer, Pará, and in the metropolitan region of Macapá, Amapá, respectively. Among 12 blood loci, alleles considered as markers of African ancestry, such as HBB*S, HBB*C, TF*D1, HP*2M, ABO*B, RH*D-, and CA2*2 were found at frequencies that are expected for populations with a predominantly African origin. Estimates of interethnic admixture indicated that the degree of the African component in Curiau (74%) is higher than that of Pacoval (44%); an Amerindian contribution was not detected in Curiau. Estimated values of African ancestry fit well with the degree of isolation and mobility of the communities. Pacoval exhibited a high proportion of immigrants among the parents and grandparents of the individuals studied, whereas persons living in Curiau exhibited a low level of mobility, despite its location in the metropolitan area of Macapá city, suggesting a relatively strong barrier against the interethnic admixture in this population. In addition, analysis of genetic data in a sub-sample consisting of individuals whose parents and grandparents were born in the study site, and that probably represents the populations two generations ago, indicated that gene flow from non-black people is not a recent event in both populations.



The slave trade brought at least 53,000 black Africans to the Amazon region. They were imported directly from Africa by the General Company of Great Pará and Maranhão, private individuals, as well as indirectly through internal trade from other Brazilian regions such as Maranhão, Bahia, Pernambuco and Rio de Janeiro (Vergolino-Henry and Figueiredo, 1990). Historical records about the slave trade to the northern region indicate that most Africans were brought from Angola, Mozambique, Kenya and Tanzania, whereas a smaller number was brought from Guine-Bissau and Cape Verde. In addition, there are records of a small number of slaves from French Guyana and Surinam introduced to the region (State of Amapá and Marajó Archipelago). Some were victims of the illegal slave trade, while others came as fugitives (Curtin, 1969; Vergolino-Henry and Figueiredo, 1990).

Following the example of other Brazilian regions and South American countries, several communities named mocambos or quilombos were founded in the Amazon region by escaped slaves, particularly in the State of Pará, a point of entry of African slaves to the region. Many of these communities still remain relatively isolated, and much of their identity is preserved.

Several genetic studies have been performed in populations descended from Africans in the Amazon region of Brazil and Venezuela evaluating the level of isolation and mobility, and quantifying the gene contribution from non-blacks among them. In this study we report genetical (12 blood genetic systems), genetical-demographic (estimates of interethnic admixture) and demographic (migration) data from two populations of predominantly African ancestry in the Brazilian Amazon region.



The mocambo of Pacoval, formed by fugitive slaves from farms of the neighboring municipality of Santarém, is located in the rural area of the county of Alenquer, along the right margin of the Curuá River, in the northwestern region of the State of Pará, at about 2oS, 55oW. Blood samples from 166 individuals were collected (63% females, ages varying from seven to 86 years).

Curiau is located in the metropolitan region of Macapá, capital of the State of Amapá, northern Brazil, at about 0o, 51oW. From 1770 Curiau constituted a point of convergence of escaped slaves from the region, including slaves from Suriname and French Guyana (Muggiati, 1971). One hundred and forty-five individuals (49% males, ages varying from four to 84 years) were sampled.

Blood samples (10 ml) were collected with EDTA as an anticoagulant, and 12 genetic systems were analyzed: haptoglobin (HP), transferrin (TF), ceruloplasmin (CP), albumin (ALB), serum cholinesterase (loci 1 and 2), carbonic anhydrase (CA2), esterase D (ESD), glyoxalase (GLO), hemoglobin (HB), ABO and RH blood groups. Phenotypic determinations were made employing classical methods of horizontal and vertical electrophoresis and immunological tests that have been previously described or referenced (Santos et al., 1983; Schneider et al., 1987; Guerreiro et al., 1989).

Allele frequencies were estimated by the maximum likelihood method using the MAXLIK program developed by Reed and Schull (1968), and the degree of racial admixture was estimated through the maximum-likelihood method of Krieger et al. (1965), using the GENIOC program.



Blood polymorphisms

All phenotype and allele frequencies were in Hardy-Weinberg equilibrium in Curiau, whereas in Pacoval the HP and CP loci showed significant deviations from equilibrium (Tables I and II). Some alleles considered to be markers of African ancestry (HBB*S, HBB*C, TF*D1, HP*2M, ABO*B, RH*D-, and CA2*2) were found at frequencies that are expected for populations with a predominantly African origin. Some unusual findings such as high frequencies of alleles ABO*B and HBB*C in Curiau, and allele RH*D in both populations, which are higher than average frequencies estimated for black African populations, may be attributed to microevolutionary factors (genetic drift and/or founder effect). In addition, the presence of the allele HBB*C in the Curiau sample (0.06) may also be attributed to the contribution of people from French Guyana to the formation of this population, since among the slaves brought to this country about 74% of them were from Senegambia and Guinea in West Africa (Curtin, 1969), where the HBB*C allele is relatively common. HBB*C was not found in Pacoval, nor in Trombetas or Cametá, two other communities of African descent in the Amazon region of Brazil (Schneider et al., 1987; Bortolini et al., 1992), but polymorphic frequencies were found in Curiepe and Panaquire, two Afro-Venezuelan populations (Bortolini et al., 1992; Castro de Guerra et al., 1996), and in various urban populations from the Brazilian Amazon characterized by a white + Amerindian + African contribution (Ayres et al., 1976; Santos et al., 1983, 1996; Cayres et al., 1991). Thus, these findings indicate that the contribution of people from West Africa to the formation of the Amazonian populations was heterogeneous.

Interethnic admixture

Estimates of interethnic admixture obtained for the two populations (Table III) indicate that the degree of black component in Curiau is significantly higher than in Pacoval. These results fit well with the apparent morphological classification of the individuals sample, which indicated 14% individuals with mixed ancestry in Curiau, and 49% in Pacoval. Interethnic admixture values are also in accordance with the level of isolation and mobility of the two populations. The proportion of immigrants among the parents and grandparents of the individuals sampled in Pacoval is high, whereas Curiau, despite its location in the metropolitan area of Macapá city, exhibits a low level of mobility (Table IV), suggesting a relatively strong barrier against interethnic admixture in this population. The proportion of African ancestry obtained for Curiau is the highest reported among communities composed of descendants of black Africans thus far studied in the Amazon region (Table III). In addition, Curiau is the first Amazonian population where an Amerindian contribution was not detected.

In order to evaluate whether admixture with Caucasians and Amerindians has occurred in the last two generations, a subsample of subjects whose parents and grandparents were born in the study site was analyzed. There where no significant differences (Tables I and II) with respect to the total sample (Pacoval: s_q2.gif (1030 bytes) = 5.95, d.f. = 12, P = 0.91; Curiau:s_q2.gif (1030 bytes) = 1.00, d.f. = 12, P = 1.00), indicating that admixture with non-blacks is not a recent event in both communities.

In summary, the genetic variability observed in Pacoval and Curiau is consistent with the African origin of the populations, with some degrees of local differentiation and admixture with people of Caucasian and/or Amerindian ancestry. In addition, the presence of people from West Africa among the African slaves brought to the Amazon region was well characterized by the occurrence of the allele HBB*C in Curiau.



A região amazônica brasileira abriga diversas comunidades formadas por descendentes de escravos africanos, algumas das quais ainda relativamente isoladas. Nós estudamos duas dessas comunidades (Pacoval e Curiau), localizadas na área rural do município de Alenquer, Pará, e na região metropolitana de Macapá, Amapá, respectivamente. Entre os 12 loci investigados, alelos considerados como marcadores de ancestralidade africana, tais como HBB*S, HBB*C, TF*D1, HP*2M, ABO*B, RH*D- e CA2*2, foram encontrados com freqüências elevadas, de acordo com o esperado para populações com origem predominantemente africana. As estimativas de mistura racial indicaram que o percentual de contribuição africana em Curiau (74%) é maior do que em Pacoval (44%) e que em Curiau não foi detectada mistura com indígenas. As estimativas de ancestralidade africana estão de acordo com o grau de isolamento e com a mobilidade das comunidade estudadas. Pacoval apresenta um elevado percentual de imigrantes entre os pais e avós dos individuos estudados, enquanto que a comunidade de Curiau, apesar de situada na região metropolitana de Macapá, apresenta uma pequena proporção de imigrantes, sugerindo a existência de uma barreira relativamente forte contra a mistura interétnica. A análise dos dados genéticos foi também efetuada em sub-amostras constituídas apenas por indivíduos cujos pais e avós nasceram na localidade em estudo e que provavelmente representam essas populações duas gerações atrás. Os resultados indicaram que a mistura com não-negros nas duas populações não é um evento recente.



We are grateful to the communities studied for their collaboration, and to Roberto A.N. Madeiro, Vilson C. Monteiro, Edson S. Abrahim Filho and Hailton O. Monteiro for assistance with the blood sample collection. This work was supported by UFPA and CNPq.



Ayres, M., Salzano, F.M., Franco, M.H.L.P. and Barros, R.M.S. (1976). The association of blood groups, ABH secretion, haptoglobins and hemoglobins with filariosis. Hum. Hered. 26: 105-109.         [ Links ]

Bortolini, M.C., Weimer, T.A., Franco, M.H.L.P., Salzano, F.M., Layrisse, Z., Schneider, H., Schneider, M.P.C. and Harada, M.L. (1992). Genetic studies in three South American Black populations. Gene Geogr. 6: 1-16.         [ Links ]

Bortolini, M.C., Weimer, T.A., Salzano, F.M., Callegari-Jacques, S.M., Schneider H, Layrisse, Z. and Bonatto, S.L. (1995). Evolutionary relationships between Black South American and African populations. Hum. Biol. 67: 547-559.         [ Links ]

Castro de Guerra, D., Arvelo, H., Rodriguez-Larralde, A. and Salzano, F.M. (1996). Genetic studies in Panaquire, a Venezuelan population. Hum Hered. 46: 323-328.         [ Links ]

Cayres, I.M.V., Vallinoto, A.C.R., Ribeiro dos Santos, A.K.C., Santos, E.J.M., Guerreiro, J.F. and Santos, S.E.B. (1991). Variabilidade genética em populações humanas da Amazônia. Braz. J. Genet. 14 (Suppl.): 141 (Abstract).         [ Links ]

Curtin, P.D. (1969). The Atlantic Slave Trade: A Census. University of Wisconsin Press, Madison.         [ Links ]

Guerreiro, J.F., Aguiar, G.F.S. and Santos, S.E.B. (1989). Serum cholinesterase polymorphism (CHE1 and CHE2 loci) among several Indian groups from Amazon region of Brazil, and segregation of the C5 variant in families. Gene Geogr. 3: 11-20.         [ Links ]

Krieger, H., Morton, N.E., Mi, M.P., Azevedo, E., Freire-Maia, N. and Yasuda, N. (1965). Racial admixture in north-eastern Brazil. Ann. Hum. Genet. 29: 113-125.         [ Links ]

Long, J.C. (1991). The genetic structure of admixed populations. Genetics 127: 416-428.         [ Links ]

Muggiati, A. (1971). O ultimo quilombo. Rev. Manchete 9: 648-657.         [ Links ]

Reed, T.E. and Schull, W.J. (1968). A general maximum likelihood method estimation program. Am. J. Hum. Genet. 20: 579-580.         [ Links ]

Santos, S.E.B., Salzano, F.M., Franco, M.H.L.P. and Freitas, M.J.M. (1983). Mobility, genetic markers, susceptibility to malaria and race mixture in Manaus, Brasil. J. Hum. Evol. 12: 373-381.         [ Links ]

Santos, E.J.M., Ribeiro dos Santos, A.K.C., Guerreiro, J.F., Aguiar, G.F.S. and Santos, S.E.B. (1996). Migration and ethnic change in an admixed population from the Amazon region (Santarém, Pará). Braz. J. Genet. 19: 511-515.         [ Links ]

Schneider, H., Guerreiro, J.F., Santos, S.E.B., Weimer, T.A., Schneider, M.P.C. and Salzano, F.M. (1987). Isolate breakdown in Amazonia: the blacks of the Trombetas River. Braz. J. Genet. 10: 565-574.         [ Links ]

Vergolino-Henry, A. and Figueiredo, N. (1990). A presença africana na Amazônia colonial: uma notícia histórica. Arq. Público do Pará 1: 27-66.         [ Links ]

(Received April 6, 1998)

Creative Commons License All the contents of this journal, except where otherwise noted, is licensed under a Creative Commons Attribution License