Am J Hum Genet 28:257-261, 1976

Esterase D in South American Indians M. A. MESTRINER,1 F. M. SALZANO,2 J. V. NEEL,3

AND

M. AYRES4

To date, 6,967 individuals have been studied for the esterase D polymorphism [1-10]. Its frequency varies from .88 to .90 among European and Canadian whites, .90-.94 in African blacks and Papuans, and .65-.77 among East Indians, Lapps, Japanese, and Nepalese. This paper reports the distribution of esterase D in eight South American Indian tribes and its intratribal variation among the Yanomama. SUBJECTS AND METHODS

Groups Studied 1. The Makiritare is a Carib speaking tribe. The samples reported here were collected in 1967 from the village designated A by Gershowitz et al. [11]. 2. The Yanomama speak a language not readily identified with any of the main language families of South America. Material from eight villages was available for this investigation. Their exact location is given by Ward [12]. The samples were also collected during the 1967 period of field work. 3. The Parakanan, a Tupi speaking group, are noted for their very light skin. Samples were collected in 1974 by Dr. F. L. Black, from Yale University, in a place located about 100 km northwest of Maraba (4030' S; 50030' W). 4. The Gorotire, a Cayapo subgroup, speak a Ge language. Sample collection was made in 1974 at a village located at 7044' S; 51010' W. 5. The Kraho are also a Ge group. Material was collected in 1974 at the villages of Pedra Branca and Cachoeira, which are located very near to one another, a few miles from the town of Itacaja' in northern Goia's (8010' S; 47°50' W). Since the results were very similar in both places, they were pooled for this analysis. 6. The Moro is a Zamucoan speaking tribe. Samples were collected in 1975 in the locality of Maria Auxiliadora, 5 km from Isla Margarita in Paraguay, located just opposite to the Brazilian town of Porto Murtinho (18020' S; 57°50' W). Received September 25, 1975; revised November 14, 1975. This research was supported by the Fundaao de Amparo a Pesquisa do Estado de Sio Paulo, Camara Especial de P6s-graduaqio e Pesquisa da Universidade Federal do Rio Grande do Sul, Conselho Nacional de Desenvolvimento Cientifico e Tecnologico, Coordenaqao do Aperfeiqoamento de Pessoal de Nivel Superior, Fundaqio de Amparo i Pesquisa do Estado do Rio Grande do Sul, U.S. Atomic Energy Commission, and the National Science Foundation. 1 Departamento de Genetica, Faculdade de Medicina, Universidade de Sio Paulo, Caixa Postal 301, 14100 Ribeirao Preto, SP, Brazil. 2Departamento de Genetica, Instituto de Biociencias, Universidade Federal do Rio Grande do Sul, Caixa Postal 1953, 90000 Porto Alegre, RS, Brazil. 3Department of Human Genetics, University of Michigan Medical School, Ann Arbor, Michigan, 48104. 4 Laborat6rio de Genetica, Centro de Ciencias Biologicas, Universidade Federal do Parli, 66000 Belem, PA, Brazil. 0 1976 by the American Society of Human Genetics. All rights reserved.

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258

MESTRINER ET AL.

7. The Caingang is a Ge speaking group. Material was obtained in 1975 in Guarita, Rio Grande do Sul (27°30' S; 53°50' W). 8. The Mapuche have a language classified as Araucanian. Blood samples were collected in 1974 by Dr. F. L. Black on the reservation of Pedregoso, Provincia de Malleco, Chile. The isoenzyme determinations were performed in Ribeirdo Preto on blood either a few days old or glycerolized. No consistent differences were observed between these two kinds of samples. The esterases were studied with methods described by Hopkinson et al. [1]. RESULTS AND DISCUSSION

The data from the eight tribes are summarized in table 1 and figure 1. Gene Es D1 shows frequencies varying from .36 to 1. No clear geographic dine is discerned, but a region of low prevalence seems to exist in northern Brazil, that comprises

.77 *86

.36 *56 .47

1.00

.65

FIG. 1.-The distribution of the Es D1 gene in several South American Indian tribes

ESTERASE D IN SOUTH AMERICAN INDIANS

259

TABLE 1 ESTERASE D POLYMORPHISM IN SEVERAL SOUTH AMERICAN INDIAN TRIBES PHENOTYPES

POPULATION

Makiritare Yanomama

.............

.............

Parakanan .............. Gorotire ...............

Kraho .................. Moro ..................

Caingang Mapuche

2-1

1

2

42

22

5

(61)

(32)

(7)

310

98

11

(74)

(23)

(3)

4

19

14

(11)

(51)

(38)

54 (33) 36 (25) 112

75 (46) 65 (44) 0

34 (21) 45 (31) 0

(100)

(0)

(0)

...............

32

33

9

(45) 14 (27)

(12)

...............

(43) 35 (69)

2

TOTAL

Es D1 FREQUENCY

.77

69 * 419

.86

..

37

.36

163

.56

...

.47

146 ...

112

1

...

74

.65

...

51

.82

(4)

NOTE.-X2 for heterogeneity: x2 = 324.1, 14 df, P < .001. Number in parentheses represents percentage of total number in each village.

the territory of the Parakanan, Gorotire, and Kraho. It is interesting to note that while the latter two tribes speak similar languages (Ge), the Parakanan speak Tupi and have other genetic characteristics that differentiate them from other Brazilian Indians (Salzano et al., unpublished). The absence of Es D2 among the Moro is also remarkable. This tribe also has a complete absence of Diego blood type (Salzano et al., in preparation). The marked differences observed in the frequency of the Es D1 allele among tribes suggests that this will be a particularly useful marker in a continuing effort to understand the biological relationships between Indian tribes through the use of multivariate measures of genetic distance (cf. [ 13, 14] ). Table 2 contrasts the findings in eight different Yanomama villages, all situated in the Roraima territory of Brazil. The precise location of these villages is indicated elsewhere [12]. Given the marked genetic microdifferentiation observed in the Yanomama (cf. fig. 1, [15]), the intervillage variation does not appear exceptional. Seventy percent of each village has been sampled, and since the individuals concerned are biologically related, the usual x2 test for heterogeneity is inappropriate. To some extent this applies to the contrast of tribes in table 1, but there the heterogeneity persists even if we divide the x2 by the factor of 5-10 suggested by Smith [16] under such circumstances.

MESTRINER ET AL.

260

TABLE 2 INTERVILLAGE VARIATION IN THE FREQUENCY OF Es DI AMONG THE YANOMAMA PHENOTYPES

VILLAGE

03 U+V ........

1

......

03W .................. 03 KP .................

03 LMN ............... 03 Q

03T 03 S

03 R

..................

..................

.................. ..................

Total .............

2-1

2

43

19

(68) 46

(30) 21

1 (2) 0

(69)

(31)

(0)

62

4

4

(88) 54

(6) 13

(6) 0

(81)

(19)

(0)

27

6

0

(82)

(18)

(0)

35

13

3

(69)

(25)

(6)

15

8

1

(63)

(33)

(4)

28

14

2

(64)

(32)

(4)

310

98

(74)

(23)

11 (3)

TOTAL

Es D1 FREQUENCY

63

.83

...

67

.84

...

70

.91

...

67

.90

...

33

.91

...

51

.81

...

24

.79

...

44

.80

...

419

.86

...

NOTE.-Number in parentheses represents percentage of total number in each village.

SUMMARY

Significant variation in the frequency of Esterase D isoenzymes was found in 1,070 individuals belonging to eight South American Indian tribes. The Es D1 allele shows frequencies varying from .36 to 1. A region of low prevalence of this allele seems to exist in northern Brazil, involving the Parakanan, Gorotire, and Kraho. The intratribal variation observed in eight Yanomama villages located in Brazil was not exceptional. ACKNOWLEDGMENTS Thanks are due to the FundagAo Nacional do Indio for permission to study the Indians and hospitality at the Posts, as well as to A. Perez Diez, F. L. Black, G. V. Simoes, N. A. Chagnon, Priscilla Neel, R. Moreno, and W. J. Oliver for help in the field or for supplying blood samples.

REFERENCES 1. HoPKINSON DA, MESTRINER MA, CORTNER J, HARRis H: Esterase D: a new human polymorphism. Ann Hum Genet 37:119-137, 1973 2. WELCH S: Red cell esterase D polymorphism in Gambia. Humangenetik 21:365367, 1974 3. BENDER K, FRANK R: Esterase D-Polymorphismus: Darstellung in der Hochspannungselektrophorese und Mitteilung von Allelhiiufigkeiten. Humangenetik 23:315318, 1974

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4. BENKMANN H-G, GOEDDE HW: Esterase D polymorphism: gene frequencies and family data. Humangenetik 24:325-327, 1974 5. WELCH S, LEE J: The population distribution of genetic variants of human esterase D. Humangenetik 24:329-331, 1974 6. ISHIMOTO G, KUWATA M, FUJITA H: Esterase D polymorphism in Japanese. Jpn J Hum Genet 19:157-160, 1974 7. BENKMANN H-G, GOEDDE, HW: Esterase D polymorphism in Assam by cellulose acetate electrophoresis. Humangenetik 27:343-345, 1975 8. K6STER B, LEUPOLD H, MAUFF G: Esterase D polymorphism: high-voltage agarosegel electrophoresis and distribution of phenotypes in different European populations. Humangenetik 28:75-78, 1975 9. WELCH SG: Red cell esterase D in studies of paternity cases in the United Kingdom. Vox Sang 28:366-370, 1975 10. MARTIN W, OTT A: Polymorphismus der menschlichen Erythrozyten-Esterase D. Phinotypenverteilung und Genfrequenzen in Berlin (West). Blut 30:299-301, 1975 11. GERSHOWITz H, LAYRISSE M, LAYRISSE Z, NEEL JV, BREWER C, CHAGNON N, AYREs M: Gene frequencies and microdifferentiation among the Makiritare Indians. I. Eleven blood group systems and the ABH-Le secretor traits: a note on Rh gene frequency determinations. Am J Hum Genet 22:515-525, 1970 12. WARD RH: The genetic structure of a tribal population, the Yanomama Indians. V. Comparisons of a series of genetic networks. Ann Hum Genet 36:21-43, 1972 13. NEEL JV: Diversity within and between South American Indian tribes. Isr J Med Sci 9:1216-1224, 1973 14. WARD RH, GERSHOWITz H, LAYRISSE M, NEEL JV: The genetic structure of a tribal population, the Yanomama Indians. XI. Gene frequencies for 10 blood groups and the ABH-Le secretor traits in the Yanomama and their neighbors; the uniqueness of the tribe. Am J Hum Genet 27:1-30, 1975 15. GERSHOWITZ H, LAYRISSE M, LAYRISSE Z, NEEL JV, CHAGNON N, AYREs M: The genetic structure of a tribal population, the Yanomama Indians. II. Eleven blood group systems and the ABH-Le secretor traits. Ann Hum Genet 35:261-269, 1972 16. SMITH CAB: Local fluctuations in gene frequencies. Ann Hum Genet 32:251-260, 1969

Esterase D in South American Indians.

Significant variation in the frequency of Esterase D isoenzymes was found in 1,070 individuals belonging to eight South American Indian tribes. The Es...
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