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Blood groups and H-Lea salivary secretion of Brazilian Cayapo Indians.

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Blood Groups and H-Lea Salivary Secretion of
Brazilian Cayapo Indians
F. M . SALZANO, H . GERSHOWITZ, P. C. JUNQUEIRA,
J . P. WOODALL, F. L. BLACK A N D W. HIERHOLZER
Departamrnto d e Geneticu, Universidade Federal d o Rio Grande d o S u l ,
PBrto Alegre, Brazil; Department of Human Genetics, Unibersity of
Michigan, A n n Arbor, Michigan, Cosa d r Saude Santu Lticia,
Rzo d e Juneiro, Guannbara, Brazil; u n d Department of
Epidemiology a n d Public Health, Y a l e University,
New H a v e n , Connecticut
KEY W O R D S Blood groups . Brazil Indians . Genetic taxonomy
Multidisciplinary studies . Salivary secretion.
.
ABSTRACT
Five hundred and twenty-six individuals from four populations
were studied in relation to the ABO, MNSs, P, Rh, Lutheran, Kell, Lewis, Duffy,
Kidd, Diego and I systems, as well as for the Wright antigen of blood groups.
The H-Lea salivary secretion of 406 of them was also investigated. Considering
the gene markers which show variation in South American Indians, the Cayapo
frequencies are in the middle of the distribution range for genes LMS , R1 ( C D e ) ,
R2 (cDE), PI, Jka, Di" and Se but present high values of Le and Fya and low
ones of L M S , LNS , Ro ( c D e ) and/or T ( c d e ) , LNS and R* (CDE). Unusual findings i n relation to previous studies are the high prevalence of Le (a ) persons
(which, however, could be expected since the frequency of gene Se is not too
high) and the presence of one Lu (a ) and one PPI (Tja)( - ) individuals.
Comparison of the pattern of inter-village variation in relation to these polymorphisms with those furnished by historical, demographic and morphological
data gives in general reasonable concordance, but some results are difficult to
interpret. New approaches and further multidisciplinary studies are needed to
obtain a clearer picture of the genetic relationships present among different
tribes of South American Indians. as well as to understand how polymorphisms
are created and maintained in human populations.
+
+
Although as many as 43,500 South
American Indians have already been studied in relation to the ABO system, a
series of questions are still unanswered
concerning the distribution of blood group
polymorphisms in these populations. Two
of the most general ones are: in what
ways does the range of gene frequency
variation differ in this major subgroup of
our species, as opposed to those observed
in other races? and How can this variability be related to their population structure, disease patterns and cultural characteristics? Important problems like the
average degree of heterozygosity at several loci, and interaction between systems
remain untouched.
The present communication is the fourth
in a series of the Brazilian Cayapo Indians, previous papers having dealt with the
AM.
J. PHYS.ANTHROP.,36: 4 1 7 4 2 6 .
health status of one group (Nutels, Ayres
and Salzano, '67) as well as the demography (Salzano, '71) and anthropometric
variation (Da Rocha and Salzano, '72) of
three of them. They are related to a longterm project aiming at the elucidation of
the questions posed in the first paragraph
(Salzano, '61a; Neel and Salzano, '66, '67;
Neel, '71).
MATERIALS A N D METHODS
The Cayapo Indians speak a Ge language and live at present in eight still
quite isolated, semi-independent communities in the Brazilian States of Para and
Mato Grosso. Details about their history,
exact location and demographic characteristics can be found in Salzano, '71.
Four populations have been studied in
417
418
SALZANO E T AL.
relation to their blood groups; brief information about them follows. 1. KubenKran-Kegn, location: lat. 8 " lo's., long.
52'8'W.; population: 310; pacified in
1952; blood collected i n 1965 and 1968;
2. Txuhahamae, lat. 1Oo20'S., long. 53"
5'W.; population: 190; pacified: 1953;
blood collection: 1966; 3. Mehranoti, lat.
8"40'S., long, 54" W.; population: 240;
pacified: 1958; blood collection: 1969;
4. Xihrin, lat. 5"55'S., long. 51'11'W.;
population: 150; pacified: 1954; blood collection: 1970. The most acculturated are
the Xikrin followed by the Kuben-KranKegn, while the Txukahamae and Mekranoti are the least disturbed by Neo-Brazilian influences.
Blood was collected in 10 or 15 ml Becton-Dickinson vacutainers with EDTA or
ACD; every effort was made to obtain
samples from all individuals living in the
villages at the time, with the exception of
the very young (less than 2 years of age).
Specimens were refrigerated shortly after
collection and carried by air to the laboratory. Those obtained during the first
visit to the Kuben-Kran-Kegn were tested
in P8rto Alegre and Rio de Janeiro; all
other determinations were performed in
P6rto Alegre andlor Ann Arbor. The blood
typing reagents with which tests were
performed in duplicate included anti-A,
A,, B, M, M g , M Y , N, S, s , PI, C (rh'),
C" (rh"), c (hr'), D (Rho), E (rh"),
e (hr"), K, k, Kp", Fya, Fyb, Di", Wra.
Tests with the following antigens were
done only once: Porto Alegre: anti-I; Rio:
anti-Kpb, J P , Jkb; Ann Arbor: anti-U,
V', Mi", M u + H t , PPI (Tja), f (hr), Lu",
Lub, Kpb, Jsa, Lea, Leb, Jk", Jkb, Di".
All studies were carried out in tubes with
2% washed red cell suspensions. Discrepancies in the duplicate determinations
were reconciled by repeat typings on cells
preserved in glycerol a t - 2 0 ° C or in
liquid nitrogen.
The salivas were directly collected into
glass tubes and boiled immediately for 15
to 30 minutes. They were transported
under refrigeration and kept frozen in
the laboratory until tested. The determination of the H and Le" secretor status
was done in duplicate in Parto Alegre
and Ann Arbor.
Gene frequencies related to the MNSs
blood groups were calculated according
to Mourant's ('54) method. In the Rh
system a gene counting procedure was
adopted, assuming the absence of r y
( C d E ) , r' ( c d E ) and r" (Cde) (details about
the assumptions and limitations of this
and other methods can be found in Gershowitz et al., '70). The other gene frequencies were obtained by standard gene
counting or square root calculations. In
all systems in which it was ,possible to
identify the heterozygote and both homozygotes we verified if their numbers were
those expected assuming Hardy-Weinberg
equilibrium. The deviations from the theoretical values were generally very small
and in only one case (the Duffy distribution among the Txukahamae) reached the
5% level of statistical significance. Due
to the number of comparisons made this
result may have occurred by chance alone.
Restrictions can be presented to the use
of ~2 methods and other "large sample"
statistics in these populations (Nee1 et al.,
'64; Gershowitz et al., '67). Therefore, no
attempts were made to verify statistically
the significance of intertribal variations.
RESULTS
Table 1 shows the MNSs blood group
phenotypes observed in the four populations and table 2 the corresponding gene
frequencies, which are compared with
those of two other Ge groups and of
South American Indians in general. The
largest amount of intratribal variation
occurred with genes LMS and LNS.The
Txukahamae and Mekranoti show very
similar gene frequencies in relation to the
four genetic markers. Gene LNS is not
present in the Kuben-Kran-Kegn sample.
As for the intertribal comparisons, the
Cayapo do not show much departure from
the prevalences observed i n the two other
Ge tribes for the LNS and L N s genes but
the frequency of LMS is higher and that
of LMS lower than those found among
the Caingang. Taking into consideration
the whole range of South American Indian values, the Cayapo frequencies are in
the upper half of the distribution in relation to gene LMs but in the lower third
for the other markers.
Phenotype prevalences for the Rh system are given in table 3 and the respective gene frequencies in table 4. There
is no clear pattern in the intratribal dis-
419
BLOOD GROUPS AND SECRETION OF BRAZILIAN INDIANS
TABLE 1
M N S s blood group phenotypes observed in four Cayapo Indian populations
Phenotype
Txukahamae
Mekranoti
Kuben-Kran-Kegn
Xikrin
Total
~~~~
no.
MS
MSs
Ms
MNS
MNSs
MNs
NS
NSs
Ns
Total
10
32
59
2
22
30
1
4
2
162
%
no.
%
no.
%
no.
%
6
20
36
1
14
19
1
2
1
1
28
31
1
9
14
1
31
34
1
10
10
37
35
6
8
31
7
28
26
4
6
23
2
17
20
1
18
25
2
18
22
1
20
27
1
8
92
-
-
-
2
5
91
15
_
-
8
135'
2
6
-
_
_
6
-
-
1
9
~~
no.
23
114
145
10
57
100
1
7
23
480
%
4.8
23.7
30.2
2.1
11.9
20.8
0.2
1.5
4.8
-
I Forty-six bloods were tested with anti-M. -N, and - S only, yielding the following results: MS (
10;MS(-): 11;MNS(+): 9;MNS(-): l I ; N S ( + ) : 2 ; N S ( - ) : 3 .
+ );
TABLE 2
M N S s blood group gene frequencies observed in four Cayapo Indian populations compared
with those ofother Ge groups and South American Indians in general
Gene frequencies
Population
Txukahamae
Mekranoti
Kuben-Kran-Kegn
Xikrin
Total Cayapo
Xavante ( N = 537)
Caingang ( N = 76) *
S. American Indians
Range(N=11,517)3
0.18
0.18
0.28
0.23
0.220
0.366
0.545
0.61
0.61
0.49
0.43
0.541
0.408
0.129
0.08
0.06
0.00
0.03
0.036
0.098
0.065
0.13
0.15
0.23
0.31
0.203
0.128
0.261
0.000 0.857
0.093 0.944
0.000 0.321
0.000 0.685
Nee1 et al., '64;Gershowitz et al., '67.
Fernandes et al., '57.
3 Data from 97 surveys performed by different investigators.
1
2
tribution and it is curious that the Txukahamae and Mekranoti who showed similar
gene frequencies in the MNSs blood group
now present very different prevalence figures. Since no cDe (Rho) phenotypes were
observed and at least one Indian proved
to be rr (cdelcde), the gene frequency
calculations were performed assuming the
absence of Ro ( c D e ) and presence of r
( c d e ) . It is possible, however, that gene
R" ( c D e ) may have been present. The
Cayapo values are again similar to those
observed in other Ge groups; the prevalence of R" (CDE) is very low but those
of R' ( C D e ) and R2 ( c D E ) are in the middle of the range given by previous investigations in South American Indians.
The results concerning the P, Duffy,
Kidd, Diego and Lewis blood groups, as
well as the H and Lewis secretor saliva
phenotypes are given in tables 5 and 6.
Previous results with gene P1 show an
enormous range of variation, going all
the way from 9 to 100%. We have reasons
to suppose, however, that at least part of
this variability is related to technical
problems. The anti-PI reagents are somewhat irregular in their degree of association with the quantitative antigenic differences which occur in this system. For
most of the tests we have used a potent
antiserum prepared in Ann Arbor and the
prevalences of PI( ) thus found were
much higher than the one obtained in
46 bloods collected during the first expedition to the Kuben-Kran-Kegn and
tested with commercial antisera. There
exists little variation among the Cayapo
villages in relation to this gene marker;
the total Cayapo frequency is similar to
the Xavante value, but the Caingang present a much lower prevalence.
+
420
SALZANO ET AL.
TABLE 3
R h blood g r o u p p h e n o t y p e s obserued in four C a y a p o Indian pop7tlations
Phenotype
Txukahamae
CDe(RhlRhl)
CcDe ( R h , r h )
cDE (RhzRh?)
cDEe ( R h g h )
CcDEe ( R h l R h z )
CDEe ( R h z R h l )
CCDE (RhzRhs)
CDE (RhzRhz)
ce ( r h )
Total
Mekranoti
Kuben-Kran-Kegn
no.
%
no.
%
710.
50
10
20
6
60
8
8
31
6
12
4
37
5
5
14
4
36
4
30
16
4
40
4
33
1
1
38
8
34
12
79
4
4
_
_
_
162
_
-
-1 _ 1
1
1
91
-
1
1
181
76
21
4
19
7
43
2
2
1
1
-
Xikrin
1
95
32
1
19
3
40
4
1
_
_
92
-
no.
29
1
17
3
37
4
Total
>lo.
131
23
107
25
206
17
14
1
2
526
0
,
24.9
4.4
20.3
4.7
39.2
3.2
2.7
0.2
0.4
-
TABLE 4
R h blood g r o u p gene f r e q u e n c i e s observed in f o u r C a y a p o I n d i a n p o p u l a t i o n s c o m p a r e d w i t h
those of other Ge grozips unrl S o u t h A m e n c a n Indians i?i general
Gene frequencies
Population
Txukahamae
Mekranoti
Ku ben- Kran-Kegn
Xikrin
Total C a y a p o
X a v a n t e (N = 539) 1
C a i n g a n g ( N =243) 2
S . A m e r i c a n Indians
R a n g e ( N = 12,826) 3
RL (CDe)
R? (cDE)
R Z (CDE)
Ro(cDe)
r (cde)
0.54
0.35
0.45
0.54
0.475
0.588
0.529
0.34
0.59
0.44
0.41
0.429
0.329
0.344
0.06
0.01
0.04
0.03
0.039
0.044
0.056
-
0.06
0.05
0.07
0.02
0.057
0.040
0.071
_
0.123 0.958
0.025 0.732
0.023 0.308
0.000 0.207
0.000 0.233
Nee1 et al., '64; Gershowitz et al., '67.
Salzano, '61b, '64a.
Data from 104 surveys made by different investigators; in 94 of them there was no indication of the
occurrence of gene r (cde). The evidence for the presence of this gene among the Cayapo rests in only
one r r (cdeicde) individual whose parents are putatively "pure blood." The other Rh negative result
was obtained from the blood of a White captured in h i s infancy and who has adopted the Indians' way
of living, having married and procreated in the village.
1
2
3
The Fy" frequencies also have a wide
range in South American Indians. There
is more inter-village variability here, the
Kuben-Kran-Kegn and Xikrin showing,
however, similar results. Considering the
Cayapo as a whole, they present a higher
frequency i n relation to this gene than the
Xavante or Caingang. As for Jh", the Xikrin show a somewhat low incidence; the
Cayapo average is very similar to the
Xavante one. The Di" gene frequencies
separate the four villages into two groups:
Txukahamae/Mekranoti and Kuben-KranKegn/Xikrin. There is a gradation of values among the Ge, from a low 0.17 found
in the Xavante reaching 0.21 i n the Cayapo and 0.34 in the Caingang. Gene Se
shows little variation in frequency among
villages. The Cayapo present a value similar to the Caingang, which is two times
higher than the Xavante frequency. Gene
Le is practically invariant among the three
villages from which data are available,
the Cayapo now showing a frequency more
similar to the Xavante than to the Caingang; the latter present a very low incidence of Le. Comparing the Cayapo results on these six gene markers with the
range of gene frequencies observed in
South American Indians, we see that they
generally are in the middle of the observed distributions.
In accordance with their somewhat low
Se frequency (0.71 -table 6 ) the Cayapo
show one of the highest prevalences of
Le (a + ) individuals found so far in South
American Indians (8% -table
5 ; surpassed only by the Xavante, who show
a n unusual figure of 30%). The frequencies of Le ( a - b + ) and Le ( a - b - ) ob-
421
BLOOD GROUPS AND SECRETION OF BRAZILIAN INDIANS
TABLE 5
P , Duffy, Kidd, Diego, a n d Lewis blood group, H a n d Lewis secretor phenotypes
observed in f o u r Cayapo Indian populations
Phenotype
Txukahamae
Mekranoti
Kuben-Kran-Kegn
Xikrin
no.
no.
no.
no.
127
162
76
91
112
135 1
Fy (a b - )
Fy (a b +)
Fy(a-b+)
Fy(a-b-)
Total
Fy (a + I ) *
Total
Grand Total
46
84
15
1
146
16
16
162
-
Jk(a+b-)
Jk (a b + )
Jk (a - b + )
Jk (a - b - )
Total
Jk (a + )
Total
Grand Total
-
-
Pl(+)
Total
+
+
+
Di(a+b-)
Di (a b +)
Di (a- b +)
Total
Di ( a +) 3
Total
Grand Total
+
L e (a - b + )
L e (a + b - )
Le (a - b - )
Total
H.Sec.
Total
L e a Sec.
Total
-
-
89
91
91
-
-
Tot a1
no.
%
76
92
391
480
82
81
42
12
49
36
1
48.0
44.1
7.6
0.3
135
45
46
181
86
6
6
92
176
162
28
1
367
156
159
526
21
40
31
3
95
128
180
180
-
22.1
42.1
32.6
3.2
47
86
86
21
40
31
3
95
360
518
518
-
11
59
107
177
198
515
515
6
33
61
-
-
-
-
-
-
98.1
-
-
69.5
118
161
161
67
91
91
-
43
152
152
5
24
62
91
29
91
91
84
180
180
6
35
45
86
42
92
92
109
11
42
162
62
5
24
91
95
10
30
135
52
11
23
86
318
37
119
4 74
67.1
7.8
25.1
136
149
85
91
150
166
-
371
406
91.4
114
148
68
91
125
164
-
307
403
76.2
-
-
-
-
38.4
-
-
-
1 A significantly lower value (43%) was obtained with other antisera in 46 bloods collected in the first expedition.
2 Tests with anti-Fyb not performed.
3 Individuals with this phenotype in tests performed with one or two antisera.
served in their red cells are respectively
in the lower third and middle of the distribution found in 48 other surveys conducted by different investigators and
involving 6,169 South American Indians.
Theoretically it is possible to obtain further estimates of Le using data from
these blood surveys. This was not attempted because we are not sure if the
relationships between the Lewis blood and
saliva phenotypes in these populations
are those usually found in Caucasian
populations (Gershowitz et al., '67).
Table 7 presents the results concerning
antigens which showed little or no varia-
tion in the populations studied here. They
are generally in accordance with investigations performed in other South American Indian groups (table 8); the majority
of these antigens also had a limited variation in populations of other ethnic groups
as well. The ABO results deserve a special
comment. As is indicated in table 7, the
AIB individual observed among the Mekranoti is a White man captured in his
infancy and who has adopted the Indians'
way of living. The one Al and two B individuals found are his children, born
from an 0 Indian woman. A purist would
be inclined to disregard these individuals,
422
SALZANO ET AL.
TABLE 6
P, Duffy, Kidd, Diego, H and Lewis Secretor gene frequencies observed in four Cayapo Indian
populations compared w i t h those of other Ge groups and
South American Indians in general
Gene frequencies
Population
P'
Txukahamae
Mekranoti
Kuben-Kran-Kegn
Xikrin
Total Cayapo
Xavante ( N = 539) 1
Caingang ( N = 288) 2
South American Indians
Range 3
Fy"
Se
Le
0.70
0.74
0.68
0.50
0.50
0.51
-
0.15
0.19
0.27
0.27
0.215
0.166
0.342
0.707
0.365
0.772
0.512
0.632
0.252
0.1130.723
0.0000.483
0.3651.000
0.2520.632
Di a
Jka
0.53
0.60
0.59
0.59
0.570
0.639
0.223
0.61
0.86
0.76
0.78
0.702
0.540
0.541
0.48
0.49
0.46
0.33
0.448
0.421
0.090-
0.131-
1.ooo
1.ooo
-
-
Nee1 et al., '64; Gershowitz et al., '67.
Salzano, '61b, '64a,b. The number of individuals studied was not the same for all systems.
on data from several authors, a s follows: PI: 98 surveys, 13,499 individuals tested; Fy": 109 and 12,891;
/ha: 78 and 9,347; Di": 135 and 14,224; Se: 6 and 1,374; Le: 7 and 1,285.
1
2
3 Based
TABLE I
Antigensfrom eight blood group systems which showed little or no variation in
four Cayapo Indian populations
Population
System
Type
ABO
0
A1
U(+)2
VW(-)
M"-)
Mi" ( - )
MV(-)
+
P
Rh
Kell
Mekranoti
Kuben-Kran-Kegn
162
87
1
2
1
181
-
B
A1B 1
MNSs
Txukahamae
Mu Ht ( - )
PPl (Tj") ( + ) 3
PP1 (Tj 9 ( - )
Cw (rhw) ( - ) 4
f (hr) ( - ) 5
f (hr) ( +)
K(-)
k(+)
Kpa(-)
Kpb(+)
Js"( - )
Lutheran Lu (a - b + )
Lu ( a + b + )
Wright
Wrk2(- )
I
I(+)
34
49
162
49
162
-
-
135
135
181
135
135
135
-
12
1
162
17
3
162
50
162
50
-
-
-
51
181
91
-
48
1
162
162
Xikrin
Total
92
522
1
2
1
169
270
429
184
449
135
24
1
486
52
3
526
271
429
271
22 1
269
1
435
171
86
86
61
12
92
35
-
-
181
181
181
181
135
135
92
86
86
86
86
86
181
9
-
-
92
1 The person whose blood had this reaction is a White captured in h i s infancy and who has adopted the Indians'
way of living. The one A1 and two B individuals are his children born by a n 0 Indian woman,
2 Among the Txukahamae only the S ( - ) bloods were tested for U.
3 Of the 12 PPI (Tj ") ( + ) nine were PI ( - ) and three P1 (
).
4 Among the Mekranoti only C (rh') (
) bloods were tested for this antigen.
5 Only CcDEe (RhlRhz) bloods were tested for this antigen.
+
including only "putatively pure blood'
persons. We have not done this for the
following reasons: 1. The A I B man is
culturally as Indian as any other member
+
of the tribe, having in fact led punitive
expeditions against other White persons;
2. Such events probably occurred repeatedly in the past, involving children of
423
BLOOD GROUPS AND SECRETION OF BRAZILIAN INDIANS
TABLE 8
Previous studies in South American Indians with antigens showing limited variation
Gene or
phenotype
System
ABO
10
IA
IB
MNSs
UJ+'
v
(-)
VW(+)
Mg(-)
Mi" ( - )
Mi" ( + )
MV(->
P
Rh
Kell
+
MU Ht ( - )
PPI (Tja) ( +)
Cw (rhw) ( -)
k
KP"(-)
Kpb ( + )
Js"(-)
Lutheran
Wright
I
1
Js" ( + )
Lu (a - )
Lu (a +)
Wr" ( - )
Wr" ( + )
I(+)
1 (-)
No. of individuals
tested
No. of
surveys
43,671
43,671
43,671
500
6,150
1
538
5,143
220
220
220
2
48
48
1
47
47
5
282
345
198
1,108
17,329
963
1,623
1,392
8
3,788
6
5,724
3
1,691
6
Range
0.797-1.OOO
0.000-0.203
0.000-0.102
-
-
-
1
1
2
11
101
3
7
10
10
24
24
27
27
1
1
-
0.952-1.000
-
-
1
Pooled data from several authors.
other tribes and ethnic groups. The detection of these genes has historical interest but there is no other reason for isolating them from the rest of the gene pool.
As a matter of fact, they have to be considered in any effort to understand the
factors which affect the genetic variability
of these populations.
Two somewhat unusual findings recorded in table 7 are the presence of one
Lutheran ( a + ) and one PPI (Tj") ( - )
among the Txukahamae. In 24 other
surveys conducted in South American Indians involving 3,794 persons, only six
were found to be Lu (a +) - four Quechuas from Hacienda Vicos, Peru (Allen,
'59), one Paraujano from the Sinamaica
lagoon, Venezuela (Layrisse, Layrisse and
Wilbert, 'SO), and one Aymara from Santa
Fe, Bolivia (Matson, Swanson and Robinson, '66). The usual explanation for these
findings is admixture with non-Indian
persons. The isolation of the Txukahamae
makes this hypothesis unlikely but an
event like the capture of the AIB man
mentioned above could have occurred in
the past. In only two other surveys PPI
(Tja) tests have been performed; among
198 Brazilian and Venezuelan Indians all
proved to be PPI (Tja) ( + ) (Gershowitz
et al., '67, '70).
DISCUSSION
In what way can the Cayapo frequencies be related to the known distribution
of blood group and secretor polymorphisms
in South American Indians? For seven of
the 14 gene markers studied which show
variation in these populations the prevalences are in the middle of the distribution
range (LM", RI (CDe), R2 ( c D E ) , PI,
Jk", Dia and Se), for two others the
values are high (Le: 0.51 and Fya: 0.70)
while for five they are low (LMS: 0.22;
LNs: 0.20; RD ( c D e ) and/or T ( c d e ) : 0.06;
L N S and R Z (CDE): both 0.04). Unusual
findings in relation to previous studies
are the high prevalence of Le ( a + ) persons (which however could be expected
since the frequency of gene Se is not too
high) and the presence of one Lu ( a + )
and one PPI (Tj") ( - ) individuals.
Comparing now our results with those
obtained among two other Ge tribes, we
see that in general the Cayapo figures are
more similar to the Xavante than to the
Caingang. In relation to the former the
424
SALZANO ET AL.
main difference occurs at the Se locus,
the Cayapo (Ca) showing a prevalence of
0.71 and the Xavante (X) 0.36. Other differences are not as large; the three most
marked concern Fya (Ca: 0.70; X: 0.54),
LMS (Ca: 0.22; X: 0.37), and L M s (Ca:
0.54; X : 41). The four main differences
between the Cayapo and Caingang (Cg)
are related to LMS (Ca: 0.54; Cg: 0.13),
P' (Ca: 0.57; Cg: 0.22), LMS (Ca: 0.22;
Cg: 0.54), and Le (Ca: 0.51; Cg: 0.25).
If the pattern of inter-village variation
concerning these polymorphisms is related to those revealed by historical, demographic and morphological data, some
interesting observations result. On the
basis of the available historical information (see Salzano, '71) the Txukahamae
and Mekranoti should show the lowest
degree of genetic differentiation; dissimilarities between them and the KubenKran-Kegn should be more marked, while
the Xikrin, who separated earlier from
the others, should present the highest
degree of differentiation. Detailed demographic information was not available for
the Xikrin but the data from the three
other localities showed in fact that the
Kuben-Kran-Kegn could be set apart from
the other two subgroups. On the other
hand, the anthropometric analysis (Da
Rocha and Salzano, '72j yielded some
curious results. The Kuben-Kran-Kegn and
Txukahamae presented the lowest degree
of morphological differentiation (as measured by the D2 statistic), the Txukahamae and Mekranoti intermediate and
the Kuben-Kran-Kegn and Mekranoti the
largest amount of dissimilarity.
Inspection of the blood group and secretor results indicate that as expected
the Xikrin figures deviate from those observed among the Mekranoti and Txukahamae (especially in relation to genes
L M s , LNs, Jh" and Di") but not from
those observed among the Kuben-KranKegn (the only difference higher than
10% found among them occurred in relation to the prevalences of gene Jh"). On
the basis of blood group and secretor data
alone the Kuben-Kran-Kegn cannot be
clearly separated from the Mekranoti and
Txukahamae. The latter show very similar frequencies in the MNSs and Diego
systems but this is not true for the Rh
blood groups.
The distribution of the frequencies in
the 14 genes showing variation in South
American Indians and studied here shows
ranges of different orders of magnitude,
some of them very wide. A constant feature in these distributions is that despite
the fact that they have different shapes
the values do not show distinct discontinuities. Therefore, despite the vast array of variation found, no classification
of the populations studied according to
their genetic homology is possible at the
moment. Fitch and Nee1 ('69), using three
different methods, have studied the relationships of 12 Indian tribes of Central
and South America using data from seven
polymorphic systems, which were considered simultaneously. Analyses such as
these will help in the establishment of a
meaningful genetic taxonomy of the South
American Indian. But for this kind of
approach it is necessary to study a population in relation to many polymorphisms
and to have concomitant data on other
variables, to check if the associations
made are the correct ones. Therefore, at
the present pace of investigation, it is unlikely that we will soon have a clear
picture of the main pattern of genetic
variation among South American Indians
on a continental scale. However, multidisciplinary studies aiming at the answer
to the questions posed in the introduction, besides furnishing the main source
of data for this approach, may also provide some significant clues for the understanding of how polymorphisms are created
and maintained in human populations.
ACKNOWLEDGMENTS
Thanks are due to F. J. da Rocha, G. V.
Simoes and M. Ayres for help in the field
work and to the FundaGBo Nacional do
Indio and Parque Nacional do Xingu
which gave permission to study the Indians and provided all facilities. Transportation to and from the villages was made
possible by Asas de Socorro. The skillful
assistance in the laboratory determinations given by G. V. Simbes, M. Ayres,
Margarete S. Mattevi, Maria C. Mallmann
and Sidia M. Callegari is gratefully acknowledged. This work is part of the Brazilian contribution to the International
Biological Programme. Our researches are
BLOOD GROUPS AND SECRETION OF BRAZILIAN INDIANS
supported by the Conselho Nacional de
Pesquisas, Conselho de Pesquisas da Universidade Federal do Rio Grande do Sul,
Coordenaciio do AperfeiGoamento do Pessoal de Nivel Superior, FundaCiio de
Amparo a Pesquisa do Estado do Rio
Grande do Sul, FundaGPo ServiGos de
Saude Publica, Pan American Health Organization, Rockefeller Foundation, U.S.
AEC AT (11-1) 1552, and the WennerGren Foundation for Anthropological Research.
LITERATURE CITED
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group phenotypes in some aboriginal Americans. Am. J. Phys. Anthrop., 17: 86.
Da Rocha, F. J., and F. M. Salzano 1972 Anthropometric studies in Brazilian Cayapo Indians. Am. J. Phys. Anthrop., 36: 95-102.
Fernandes, J. L., P. C. Junqueira, H. Kalmus,
F. Ottensooser, R. Pasqualin and P. Wishart
1957 P.T.C. thresholds, colour vision and
blood factors of Brazilian Indians. I. Kaingangs.
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Fitch, W. M., and J. V. Neel 1969 The phylogenic relationships of some Indian tribes of
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Gershowitz, H., P. C. Junqueira, F. M. Salzano
and J. V. Neel 1967 Further studies on the
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Layrisse, M., 2. Layrisse and J . Wilbert 1960
Blood group antigens among the Paraujano.
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Matson, G. A., J . Swanson and A. Robinson 1966
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Mourant, A. E. 1954 The Distribution of the
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Neel, J. V. 1971 Genetic aspects of the ecology
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1967 Further studies on the Xavante
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19: 554-574.
Neel, J. V., F. M. Salzano, P. C. Junqueira, F.
Keiter and D. Maybury-Lewis 1964 Studies
on the Xavante Indians of the Brazilian Mato
Grosso. Am. J. Hum. Genet., 16: 52-140.
Nutels, N., M. Ayres and F. M. Salzano 1967
Tuberculin reactions, x-ray and bacteriological
studies in the Cayapo Indians of Brazil. Tubercle, 48: 195-200.
Salzano, F. M. 1961a Studies on the Caingang
Indians. I. Demography. Hum. Biol., 33: 11C130.
1961b Studies on the Caingang Indians.
111. Blood groups. Am. J. Phys. Anthrop., 19:
391404.
1964a Blood groups of Indians from
Santa Catarina, Brazil. Am. J. Phys. Anthrop.,
22: 91-106.
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from Santa Catarina, Brazil. Am. J. Hum.
Genet., 16: 301-310.
1971 Demographic and genetic interrelationships among the Cayapo Indians of
Brazil. Social Biol., 18: 148-157.
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