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Blood groups of Brazilian Indians.

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BLOOD GROUPS O F BR,AZILIAN INDIANS
A. M. P A N T I N AND P. C. JUNQUEIRA
Department of Pathology, Cambridge University, and Znstituto Oswaldo Cruz,
Rio de Janeiro
This paper reports a study of the blood-groups of 73
Brazilian Indians. The region in which the blood samples
were collected is that bounded by the rivers Batovi, Ronuro,
Kurisevu and Kuluene, which are tributaries of the Xing6 in
the Matto Grosso. The4ribes of this region are the following:
Bakairi, Kalapalo, Nahukwi, Kuikuru and Matipuhy of the
Cariba family; W a u r i and Mehinaku of the Aruaque family;
Auety and Kamaiur6 of the Tupi family; and the isolated
group Trumai.
Of these tribes, only two were accessible enough to provide
blood samples which could reach the laboratory in suitable
condition for testing. These were the Kamaiuras living near
Camp Jacar6 of the Fundacjio Brasil-Central, and the Kalapalos living in a fishing village near Camp Kuluene on the
Kuluene river.
From Camp Jacar6 we collected 35 blood samples, comprising 32 KamaiurA (Tupi), 1 Trumai, 1 W a u r i (Aruaque) and
* This work was made possible by the generosity of the Brazilian Government,
through Dr. Olympio d a Fonseca, Director of the Instituto Oswaldo Cruz; Dr.
Walter Oswaldo CNZ, Director of the Department of Haematology of that Institute; Dr. Modesto Donatini Dias d a Cruz, Director of the ServiGo de ProteGLo dos
Indios; General Borges Fortes, Presidente da FundaQLo Brasil-Central; Orlando
VilasbBas, Chefe d a ExpediqHo Xingd-Roncador, and Dr. Pedro E. Lima. To all
these, and t o the Brazilian Air Force which supplied us with air-transport, we offer
sincere thanks. We are especially indebted to Dr. A. E. Mourant, Director of the
Blood Group Reference Laboratory, the Lister Institute, who not only provided the
very valuable sera for our tests but calculated the gene frequencies from our results, and whom we would gratefully thank for his help.
395
396
A . M . PANTIN A N D P. C . JUNQUEIRA
1 Sui& (GB). The first tests on these samples were set up in
Rio d e Janeiro 28 hours after bleeding, and completed two
days later.
From Camp Kuluene, 104 samples were taken. Storms delayed our air-transport, and only 38 samples reached Rio in
first-class condition for tests. These included 24 Kalapalos,
3 Nahukw&, 2 Matipuhy, and interbreeds as follows: 4 Kalapalo/Kuikuru, 2 Kalapalo/Nahukwb, 1 Kalapalo/Matipuhy,
1 Nahukwb/Matipuhy, and 1 Nahukwb/Kuikuru. All belong to
the Cariba family. One sample only was from a cross between
different families, viz. Kalapalo (Cariba) /Mehinaku (Aruaque). This second batch of tests was set up three days after
bleeding and finished two days later.
MATERIAL AND METHODS
1. CeZEs. These Indians would not permit venepuncture, so
finger-puncture was used. Blood from deep finger-stabs was
allowed to drop into dry sterile tubes which were closed with
sterile rubber bungs and placed upright in sealed tins immersed in cool water (about 10.C.) in thermos containers.
These containers were transported by air from the Xingfi t o
the laboratory in Rio de Janeiro.
F o r tests, only bright firm blood-clots in clear colorless
serum were used. All' clots not absolutely first-class were discarded. Each clot was shaken up in a little sterile 0.9% saline,
the loose cells removed and washed, and an approximately
2% cell suspension, as judged by eye, reconstituted in sterile
saline.
2. Sera. The sera used in these experiments were supplied
by the kindness of Dr. A. E. Mourant, of the Blood Group
Reference Laboratory, the Lister Institute. They were freezedried, and only made up as needed in sterile distilled water.
They were specific for the antigens A, A,, B, C, c, D, E, e, M, N,
S, P, Lutheran, Kell, and D~iffy(FY). All except anti-hll and
anti-N, which were animal sera, were human sera of high
BLOOD GROUPS O F BR.AZILIAN I N D I A N S
397
titer, all except anti-Kell were absorbed to remove the
natural agglutinins, and all except anti-Kell and anti-Fya
were saline-aggl-utinating. Tests for Kell and Fy" were done
by the Coombs test (adding rabbit anti-human-globulin serum
to the sensitized cells) and by the direct action of the testing
serum on cells suspended in 20% human albumin. All the sera
were used undiluted, except anti-A and anti-B, which were
diluted to half-strength, lest these very powerful sera should
lyse the cells.
3. Methods. The small-volume technique devised by Race
was employed throughout, whereby equal volumes (approuimately 0.008 ml) of serum and of 2% cell-suspension in saline
are delivered from a marked Pasteur pipette, under direct
vision, into clean d r y precipitin tubes and kept for the necessary time at the appropriate temperature. Tests for A, A,, and
B were kept for two hours at room temperature. Tests for Rh
antigens were incubated for 90 minutes at 37"C., and never
centrifuged. Larger volumes were used in the tests f o r M and
N. These, and tests for S, P and Lutheran were kept f o r two
hours in a refrigerator at approximately 18°C. F o r Kell. and
Fy", 1 volume of 10% cell-suspension was incubated with 2
volumes of serum for 30 minutes at 37"C., and the cells, after
3 washings in saline, exposed to rabbit anti-human globulin
Anti-Kell was also
serum (titer $3, no zoning; used a t -io-).
used in equal volumes with a 2% suspension of cellms in 20%
human albumin. When the test was ready, the contents of each
tube were placed on a slide with minimal disturbance and read
under the microscope (Q" objective). Controls of known cells
were set u p with each test. All the results were viewed by one
person.
Further comments on the tests will be made under the separate headings. The gene-frequencies were calcul-ated from our
data by Dr. A. E. Mourant, using his method which is described
in his papers on Basques and Indians (Chalmers et al., '49;
Prasad et al., '49).
398
A . M . PANTIN AND P. C. JUNQUEIRA
RESULTS
The ABO groups
The cells only, aiid not the serum, of the blood samples were
examined. All these Brazilian Indians belonged to Group 0.
Of 27 White controls, the ABO groups were :
GROUP
N O . OBSERVED
FREQUESCY
4
4
0.2962
1
1
0.0741
3
14
0.1111
0.5186
37
1.0000
The “ Rhesus” Blood groups
The sera used in these tests were of high titer and absolute
specificity, aiid were used undiluted; the resu1,ts were completely definite in all cases. An acute shortage of anti-e made
it impossible to test all, or many, of the bloods for this antigen,
and such tests as were made were disregarded in calculating
chromosome frequencies. Apparently little information is thus
lost. The phenotype ccddee ( r r ) was not observed in these
Indians.
RE.4CTIONY
anti C D E c e
++++
++-++-+
-++++++-+-+
XO. OBSERVED
I,HENOTYPE
COMPIOXEST
GEPI’OTYPE
CeDE
CCDee
CeDee
ccDEE
CCDE
ccDce
Ri Rz
17
R, Ri
Ri R o
R3 R,
11
Ri R‘
R, R,
Tupi
TOTAL
32
23
6
0
4
15
11
3
6
0
0
-
-
0
-
38
35
73
0
9
6
4
399
BLOOD GROUPS O F BRAZILIAN INDIANS
~
c~~~~~~
NO.
OBSEBVED
CcDE
CCDee
CcDee
ecDEE
CCDE
ecDee
NO.
FREQUENCY
BXPECTED
OBSERVED
28.14
25.06
5.63
9.30
4.56
0.32
73.00
~~~
FREQUENCY
EXPECTED
0.4384
0.3014
0.1233
0.0822
0.0548
0.0000
0.3855
0.3433
0.0770
0.1275
0.0624
0.0043
1.0001
1.0000
Chromosome frequencies:
CDe
cDE
eDe
CDE
%
R,
R.
R.
0.5859
0.2973
0.0657
0.0510
0.9999
Of 27 White control bloods, the Rh groups were as follows:
PHENOTYPE
COXNONEST GENOTYPE
NO. OBSERVED
~~
6
4
1
6
4
4
2
CCDee
CcDE
ccddEe
ceDE
ccddee
CeDee
eeDee
27
The M N S groups
The interpretation of the MNS results presents a problem.
The anti-hi and anti-N sera used in these experiments were
animal sera prepared at the Blood Group Reference Laboratory a t the Lister Institute. Two different sera of the same
specificity were used for every test, and these same sera had
been in routine use at the Lister with satisfactory results
against controls. In addition, these same sera gave the following results with 27 White European bloods used as controls in
Brazil: 11MN, 9 M, and 7 N, a distribution very close to the
expected; and these results agreed with tests made on the
400
A . M . PANTIN A N D P. C. JUNQUEIRA
same control bloods with commercial sera. The physical conditions of the tests were rigidly controlled. Nonetheless, the
Brazilian Indian bloods gave a number of N-positives which
were neither in accord with previous findings of other workers,
nor statistically likely, and this feature recurred in the first 19
of a series of North American Indian bloods (Dieguefios)
tested with the same sera. When a different pair of anti-N
sera was employed to test the remaining Diegueiios, the numbers of N-positives dropped. On the other hand, a re-test of
N-negative Brazilian Indians with the original sera still gave
a negative resu1.t.
There a r e therefore three possibilities :
1. That the first pair of anti-N sera, which a r e notoriously
difficult to make, contained non-specific agglutinins and gave
a number of false positives.
2. That these sera contained in addition to anti-N an agglutinin against some unknown blood antigen which was picked
up in different proportions of different populmations.
3. That the high incidence of N in the peoples tested is real.
I n view of the reaction with the controls, and the fact that MM
Indian bloods still gave a negative reaction on re-test with the
doubtful anti-N sera, this is at least a possibility.
Without further experiment, it is impossible to determine
which is the right answer, and the matter must be left there
for the present.
The anti-M serum, and the human anti-S serum gave excellently clear-cut results.
Indians: MNS group
PHENOTYPE
NO. OBSERVED
MS
Mas
MNS
MNss
NS
NSS
8
6
35
34
0
1
74 '
One additional sample was tested f o r MNS, making 74.
401
BLOOD GROUPS O F BRAZILIAN I N D I A N S
Probably only one N, the rest being MN
NO. OBSERVED
M+
M-
FREQUENCY
73
0.9865
0.0135
1
-
-
74
1.0000
+ M.
GENE FREQUENCIES
M
N
0.8837
0.1163
1.0000
The evidence is insufficient t o show how S genes are distributed between M and N.
Results of the White control tests were :
PHENOTYPE
NO. OBSERVED
FREQUENCY
MS
4
MI38
MNS
MNss
5
6
5
NS
1
Nss
-6
0.1482
0.1852
0.2222
0.1852
0.0370
0.2222
Total:
27
1.0000
Th.e Lu t hmwn antigen
This antigen was discovered in 1946 by Callender and Race
('as), who found its antibody in the serum of a man who had
been many times transfused. This antibody has since been
found, always a s an immune response, on 5 further occasions.
The Lu" antigen is inherited a s a Mendelian dominant character (Callender and Race, '46) and the phenotypes Lua+ and
Lua- can be recognized by the anti-Lutheran serum. Race and
Sanger ('50) give the incidence of the Lutheran group in
Engl'ish blood as follows :
Total no.
LU.+
105 (7.65%)
1373
Lu'1268 (92.35%)
And the gene frequencies :
Lub = 0.9610
and Lila = 0.0390
1.0000
Mainwaring ancl Pickles ( '48) describe strongly and weakly
reacting types of cells, and deduce the existence of three allelomorphic genes determining the nature of the Lu" substance.
402
A . M . P A N T I N A N D P. C. JUNQUEIRA
I n the Brazilian Indians in the present study, 3 of the 12
positive results were weak, 9 strong.
Indians :
PHENOTYPE
NO. OBSERVED
FREQUENCY
LU.+
Lu'-
12
61
0.1644
-
0.8356
73
1.0000
Gene frequencies :
Lub = 0.9141
Lu' = 0.0859
1.0000
Controls :
NO. OBSERVED
LU*+
Lu'-
FREQUENCY
2
0.0740
0.9260
37
25
1.0000
The P untigeta
A single human anti-P serum was used, and the test carried
out at temperatures between 10°C. and 18°C. Positive results
graded from large coarse agglutinates comprising all cells to
weak agglutinates of a few cells among many unagglutinated
ones. Four categories were established (#,
[
and weak)
and a positive was recorded if any signs of general agglutination were present. But it was not always easy to interpret the
results with this serum, as the border-line between a very weak
positive and a negative was difficult to determine.
+, +]
~
PHENOTYPES
NO. ORSERVED
FREQUENCY
P+
P-
30
43
0.4110
0.5890
73
1.0000
~
Indians :
Gene frequencies:
~-
p = 0.7675
P = 0.2335
Controls:
PHENOTYPES
N O . OBBERVED
EREQUENCY
P+
14
13
0.5185
0.4915
P-
I
27
1.0000
403
BLOOD GROUPS OF B R A Z I L I A N I N D I A N S
T h e Kell antige9z
I n 1946, the Kell antigen was identified by Coombs, Mourant
and Race. Anti-Kell, in the “incomplete” form, had developed
in the serum of a Kell-negative mother whose child was
thought to have haemolytic disease. The Kell antigen was
found to be inherited as a Mendelian dominant character.
Race and Sanger ( ’50) give the frequency of the Kell antigen in Caucasian bloods (White American and English) as
f OllO\VS :
Kell+
Kell-
NO. OBRBRVED
FREQUENCY
43
380
0.1017
0.8983
423
1.0000
-
and the gene frequencies
k = 0.9478
K = 0.0522
Dunsford (’49), testing 566 Group-0 bloods with a different
anti-Kell serum, found the percentage of K positives to be 7.24,
while Bertinshaw e t al. ( ’50), using the same serum on a
further random sample of 475 bloods, found 6.95% were E
positive. The antibody in the anti-Kell serum used to test
these Brazilian Indian bloods was “incomplete,” and the
serum, Group 0, was not absorbed; as the Indians were all
Group 0, this was immaterial.
PHEIOTYPES
h0. OBBERVED
FEEQUENCY
K’
K-
17
56
0.2329
0.7671
73
1.0000
Indians :
Gene frequencies:
k = 0.8759
= 0.1241
K
1.0000
Controls :
PHEPFOTYPES
NO. OBSERVED
FREQUENCY
K+
2
25
0.0740
0.9260
27
1.0000
K-
-
404
A . M . PANTIN A N D P. C. JUNQUEIRA
The "D26fy" antigen, Fy"
The most interesting thing about these Indian bloods is that
they all appear to lack the antigen Fy".
The example of anti-Fy" used t o test these Indian bloods did
not agglutinate cells in saline, and the reaction on Fy-positive
cells in 20% albumin was either extremely weak o r negative.
Each test was therefore done by the Coombs method. All 73
Brazilian Indian bloods gave negative results. The same serum
used by the same method gave strong positives with 8 out of
16 White control, bloods (Portuguese, English, French and
German).
The blood-group antigen Duffy, o r Fy", was discovered by
Cutbush, Mollison, and Parkin ( '50), who found the corresponding antibody in the blood of a man immunized by transfusion. The most complete published figures, those of Race and
Sanger ( '50),show that Fy" is present in the blood of 65% of
English people. Cutbush and Mollison ('50) have found it in
91% of persons from Northern Pakistan; Miller, Rosenfield
and Vogel ( '51) in 99% of Chinese and 26% of American Negroes ; and Hartmann and Brendemoen in 97% of Norwegian
Lapps. It is a n interesting fact, which may prove to be of some
anthropological importance, that the antigen Fy" has not been
found in this group of Brazihian Indians.
SUMMARY
Blood from 73 pure-bred Brazilian Indians from the hlatto
Grosso (Kalapalos and Kamaiuras) has been examined for the
antigens A, A,, B, C, c, D, E, M, N, S, P, Lutheran, Kell, and
DUffY (FYI.
All these Indians belonged to group 0. All carried the antigen D. The phenotype ccddee (rr) was not observed. An unexpectedly high number of N-positives was found. Whether
this finding was real or due to non-specific agglutination is
discussed. Of these Indians, 58.9% were P-negative. The most
interesting fact was that all the Indians tested were Duffynegative. This fact may prove to be of anthropological8importance.
a
Personal cominunication.
BLOOD GROUPS O F BRAZILIAN INDIANS
405
L I T E R A T U R E CITED
BERTINSHAW,
DOREEN,S. D. LAWLER,H. A. HOLT, B. H. KIRMANAND R. R. RACE
1950 The combination of blood-groups in a sample of 475 people i n a
London hospital. Ann. Eugen., 15: 234-242.
CALLENDER,SHEILA
T., AND R. R. RACE 1946 A serological and genetical study
of multiple antibodies formed in response t o blood transfusion by a
patient with lupus erythematosus diffusus. Ann. Eugen., 13' : 102-117.
CHALMERS,
J. N. M., E. W. IKINAND A. E. MOURANT 1949 The ABO, MN, and
Rh blood groups of the Basque people. Am. J. Phys. Anthrop., n.8. 7:
529-544.
COOMBS,R. R. A., A. E. MOURANTAND R. R. RACE 1946 In-vivo isosensitization
of red cells in babies with haemolytic disease. Lancet., i: 264-266.
CUTBUSH,MARIE, AND P. L. MOLLISON 1950 The Duffy blood group system.
Heredity, 4: 383-389.
CUTBUSH,MARIE,P . L. MOLLISONAND D. M. PARKIN1950 A new human blood
group. Nature, 165: 188.
DUNSFORD,I. 1949 A saline-agglutinating Kell antibody. Nature, 163 : 962.
MAINWARINQ,
U. R., AND M. M. PICKLES1948 A further case of anti-Lutheran
immunisation, with some studies on its capacity f o r human sensitization.
J. Clin. Path., 1 : 292-294.
MILLER,E. B., R. E. ROSENFIELD
AND P. VOQEL 1951 On the incidence of some of
the new blood agglutinogens in Chinese and Negroes. Am. J. Phys.
Anthrop., n.s. 9 : 115-126.
PRASAD,
C. H., E. W. IKINA N D A. E. MOURANT 1949 The Rh and M N S blood
groups of some students from India. Am. J. Phys. Anthrop., n.s. 7:
553-558.
RACE,R. R., AND R. SANQEE1950 Blood Groups in Man. Blackwell Scientific
Publications, Oxford.
POLYNESIAN-AMERIND
RELATIONSHIP.The blood grouping figures
favor neither Melanesia nor Micronesia [as the probable path of the
ancestors of the Polynesian]. If no other facts were known they would
almost compel one to look for another alternative. And the only obvious alternative, the very long landless ocean path from America,
gains strong support from blood gene frequencies.
The serological similarity of these two groups which is not unsupported by likenesses in other criteria suggests that the Polynesian
406
AMERICAN ASSOCIATION O F P H Y S I C A L ANTHROPOLOGISTS
and the Amerind have a coinmon dominant. component. Further, this
component, perhaps somewhat differentiated during centuries in the
widely diverse environments of the Americas, could have reached
Oceania a t any time with the aid of prevailing ocean currents. That
ocean currents have had a major influence on the migratory journeyings of the Polynesian has gained much support in recent years
following the successful enterprise of Heyerdahl and the involuntary
voyages of two other groups quoted by Avias, 1949.
These three remarkable ocean voyages took place within a space
of 12 months. How many similar sailings have taken place over the
centuries we have no idea, but we know that long ocean voyages taking
advantage of ocean currents and prevailing winds were well within
the capabilities of the seafaring early Polynesian.
Avias ( ’49) has suggested that the Polynesian has resulted from
admixture of an Amerind component with a proto-Melanesian group
which had arived in Polynesia before the Amerinds. H e described the
latter group as a negrito-ainoid complex with possibly a dash of
Papuan. Our results suggest that the proto-Melanesian component
must be weak in Polynesia, as it has left little serological evidence
of its presence in the Maoris.
Nevertheless there appears to be much to indicate that another
race reached some of the islands of Polynesia.
This group, though perhaps outnumbered by the “Amerinds,” may
well have been responsible for some of the similarities of culture and
language that have been quoted in support of an eastward course for
the Polynesian. It is also likely that since his original settlement in
the central Pacific the Polynesian has, on occasion, driven westward
into both Melanesian and Micronesian territories and possibly even
into Indonesia.
In conclusion it is submitted that the serological evidence presented
in this paper strongly supports a Polynesian-Amerind relationship,
making it probable that the islands of Polynesia have been settled
largely by migrations f rom continental America. Prevailing ocean
currents and other factors suggest the coast of Peru as the starting
point of such migrations.- J. J. Graydon. Blood groups and the
Polynesians. lIankind, vol. 4, no. 8, March, 1952, pp. 329-339.
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