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Blood group serum protein and red cell enzyme groups of Amerindian populations in Colombia.

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Blood Group, Serum Protein and Red Cell Enzyme Groups of
Amerindian Populations in Colombia
R. L. KIRK,' E. M. McDERMID,' N. M. BLAKE,' D. C. GAJDUSEK,2
W. C. LEYSHON AND R. MAcLENNAN
1 Department of H u m a n Biology, T h e J o h n Curtin School of Medical Research, Canberra, Australia; National Institute of Neurological Diseases
and Stroke, National Institutes of Health, Bethesda, Maryland; 3 H u m a n
Genetics Branch, National Institute of Dental Research, National Institutes
of Health, Bethesda, Maryland; 4 International Center for Medical Research
and Training, T u l a n e University-Universidad del Valle, Cali, Colombia.
KEY WORDS South American Indians. Colombia. Blood groups.
Serum proteins . Cell enzymes.
ABSTRACT
Red cell samples from persons belonging to four Amerindian linguistic groups in Colombia were investigated for genetic variants in eight blood
group systems: for three of the groups investigations were extended to ten red cell
enzyme and four serum protein systems. The groups studied are the Noanama
(including six Empera) of the Rio Siquirisua and Rio Docampado on the Pacific
lowlands and the Cofan, Ingano and Siona Indians of the Upper Rio Putumayo
and its tributaries to the east of the Andes. Only blood group 0 was present
among two of the groups and the same groups were 100% Kp(b +), k in the Kell
system. Di(a+) frequencies were high in three groups and there was marked
variation between groups for the MNS, Rh, P, Lewis and Duffy systems. Polymorphism in all the three linguistic groups studied for serum proteins and red
cell enzymes was present only in the red cell acid phosphatase, phosphoglucomutase (locus-1) and haptoglobin systems. 6-phosphogluconate dehydrogenase
was polymorphic in the Noanama, and caeruloplasmin was polymorphic in the
Ingano linguistic group. In addition two persons belonging to the Cofan linguistic
group revealed the presence of an "atypical" component in the lactate dehydrogenase system. No variation was found in the other six red cell enzyme and two
serum protein systems.
Comparison with published data on red cell enzyme and serum protein groups
for other South American Amerindian populations shows the Colombian populations studied here most closely resemble the Cayapo of Brazil.
Approximately 6 % of the 175 million
persons in South America are recognized
as Amerindian, and of these i t has been
estimated that about one-tenth are unacculturated at the present time (Salzano,
'68). These are distributed among a large
number of endogamous groups or tribes
differentiated frequently by geographical
location and socio-cultural patterns of
which distinctive language or dialect is
often the most striking. During the last
decade increasing attention has been
given to the genetic structure of such
groups, and a number of them particularly in Brazil, Peru and Venezuela have
been studied in some detail (Arends et al.,
'67; Arends et al., '70; Fitch and Neel,
AM. J. PHYS.ANTHROP.,41: 301-316
'69; Johnston et al., '69; Neel et al., '64;
Neel and Ward, '70; Salzano, '71, '72; Salzano et al., '72a; Salzano et al., '72b;
Shreffler and Steinberg, '67; Tashian et
al., '67; Weitkamp and Neel, '70). In addition to these investigations a large number of studies of the distribution of red cell
antigens and a smaller number of some of
the serum protein groups were summarized
up to November 1967 by Post et al. ('68)
and more recent investigations together
with new data on the red cell enzyme and
serum protein groups of the Cayapo of
Brazil are included in Salzano et al. ('72b).
Relatively few of the population genetic
studies carried out previously have involved groups living in Colombia, a n d with
301
302
KIRK, McDERMID, BLAKE, GAJDUSEK, LEYSHON AND MAcLENNAN
the exception of one study of the Ica and
Paez which included investigation of transferrins (Arends and Gallango, '65) all
were concerned with red cell antigens,
mainly the ABO blood groups. Only two
populations, the Ica and Tunebo, have
been tested for a range of red cell antigen
systems, ABO, MNS, Rh, Jk, Fy, Di, P and
Lewis (Layrisse et al., '63).
Because of the relative paucity of data
on red cell enzyme and serum protein
groups among Amerindian populations in
South America and particularly the absence of any kind of such information for
Colombian Amerindian populations we
considered it fortunate that we had the
opportunity to visit Colombia and collect
samples from persons representing several
linguistic groups.
peans. Rare marriages with the Empera
do occur (Gajdusek, '72).
Intensive anthropological work on the
Siguirisua was conducted in '64-'65 by
Elizabeth Kennedy ('72). The geography
of the area has been described by West
('57) and Eder ('63). Approximately 50%
of the Siguirisua River population are included in this study.
The Empera live mainly to the north on
the other river systems, although a small
enclave was found on small tributaries of
the Rio Docampado (fig. 2). There are
over five times as many Empera as Noanama speakers. We studied six individuals
from one household of Empera on the
Quebrada Basura, a small tributary of
the Docampado river below the Rio Siguirisua.
ETHNOGRAPHIC BACKGROUND
The Putumayo Indians
The Putumayo Indians live on the flat
forested lowlands of the Rio Putumayo. The
small Indian population is in close contact
with and is greatly outnumbered by colonizing Mestizos. Mestizo and Black settlers
live in scattered homesteads along the
rivers between the Indian enclaves. There
is very little social contact or intermarriage between the Indians and these settlers. The Indian enclaves we studied,
shown on the map in figure 3 , belong to
three groups: Cofan, Ingano and Siona
(Igualada and Castellvi, '40). We were told
of a small settlement of recently immigrant
Chocoan Indians on the Rio Acae (Gajdusek, '72). There are no other known contacts between the Choco and the Putumayo
groups. The three Putumayo groups studied marry infrequently with each other.
The offspring of marriages with mestizos
tend to move out of the Indian community.
The Cofan live along the San Miguel,
Aguarico and Guamues rivers. Their population of some 550 people extends into
Ecuador. The Cofan language has not
been found to have a n y definite relation to
any other linguistic family of American
Indian languages (Robinson, '70). Although
dialects vary on the above rivers, the culture is the same. Only the Cofan living
along the Rio San Miguel and its tributaries are included in this study.
The Ingano speaking the Quechua language of the Ingas (Incas) are scattered
over a wide area of the lowlands (Ortiz,
The Indians we have studied are from
two lowland areas of Colombia, the Pacific
and the Putumayo. Their geographic locations are indicated in the maps 'in figures
1 to 3. Nearly all Indians in the Pacific
lowlands are Chocoans, estimated by Holmer ('63) to total 10,000. They speak Carib
languages that can be divided into two distinct groups: the Empera or Northern
Choco and the Noanama, also called the
Southern Choco or Waunana (Stout, '48;
Wassen, '63). The Putumayo groups are
separated from the Chocoans by the whole
Andean cordilleras. There are no linguistic
or ethnological data suggesting relationships between the Indians of the two regions.
The Pacific Indians
The Noanama whom we studied live
along the Siguirisua River. They are relatively isolated and unacculturated and
their population of 300 is approximately
14% of the total Noanama (Reichel-Dolmatoff, '60; '62). They live in about 20
large households scattered along the length
of the Rio Siguirisua, a large tributary of
the Docampado river (fig. 2). Blacks of
African origin inhabit the lower reaches
of the Rio Docampado and the Rio Siguirisua; although they are close neighbors
with the Noanama there is little social
contact between them. There was no evidence of any intermarriage between the
Noanama and these Blacks or with Euro-
SERO-GENETIC STUDIES IN COLOMBIA
303
Pacific
Ocean
Bogota
Buenaventura
*Call
km
-
0
50
100
150
t
I - ( -
Putumavo Indians
z
ECUADOR
Fig. 1
Geographic location of the two study areas in Colombia.
-7
304
KIRK, McDERMID, BLAKE, GAJDUSEK, LEYSHON AND MAcLENNAN
......
..':...
::.::::'.,::
.............,...,...
.'...:.;f(:;.'.
...._.. ...
.......
.....................
. .. ....
km
0
Fig. 2 Distribution of the Empera and Noanama linguistic groups of Chocoan Indians on
the Pacific side of the Andean cordilleras (adapted from Reichel-Dolmatoff, '60).
305
SERO-GENETIC STUDIES IN COLOMBIA
0
2
0,
9
0
2
0,
9
M
a
;
1
0
F
3
306
KIRK, McDERMID, BLAKE, GAJDUSEK, LEYSHON A N D MAcLENNAN
'65). These Quechua speaking peoples
have reputedly descended from the Andean
Altiplano of Colombia or Ecuador, where
some groups, such as one in the Sibundoy
valley of Colombia, still speak Quechua.
However, the relation between present day
highland and lowland Quechua speakers is
not clear. We studied one large group of
Ingano speakers, who reside at San Marcelino on the San Miguel river.
The Siona live along the Putumayo river
(fig. 2). They have been fragmented since
1932 (Chaves, '45) and their present sittuation is one of cultural disintegration
(de Recasens, '64-'65). Their language is
in the Western Tukano family (Ortiz, '65)
whose speakers are scattered in Colombia
and Ecuador. There are perhaps 300 Siona
of whom some 200 live on both sides of the
Putumayo at Buena Vista.
with a range of antisera belonging to eight
red cell antigen systems. In the case of the
Siona a few of the samples were found to
be unsuitable for typing and the totals
have been adjusted appropriately. Further,
because of the small number of Empera (a
total of six from one family group) they
have been included in all the tabulations
with the Noanama. Separate evaluation of
these six individuals revealed no unusual
phenotypes in any of the systems studied.
ABO
South American Amerindian populations
are characterized by extremely high frequencies of blood group 0, achieving 100%
in many cases. The present study, as shown
in table 1, is no exception, only eight of
361 persons tested being other than group
0. These eight individuals comprised three
group A1 persons among the Ingano, and
MET H 0D S
three group A1 and two group B persons
Blood samples were taken by venepunc- among the Siona. Similar low frequencies
ture into "Bayer" venules and carried for for the A and B blood groups have been reperiods ranging from a few hours to a few ported for a number of other Amerindian
days at a temperature of 4 " to 10" C until -groups in Colombia and the data have been
placed on wet ice for transport to Washing- summarized by Post et al. ('68), and Salton, D.C. Here serum was separated and zano et a1 ('72a).
a portion of the clots and serum frozen at
MNSs
- 70" C. The frozen samples, except for
Tests were performed with anti-M, -N,
those from the Siona, were transported at
dry-ice temperature to Canberra where -S and -s as well as with anti-U. All samthey were stored in a liquid nitrogen re- ples were U( t). Table 2 demonstrates
frigerator until ready for testing for red cell marked differences between all four linguistic groups in the frequencies of the
enzymes and serum proteins.
Blood grouping was carried out in Wash- genes, but in three of the groups the most
ington on cells recovered from the un- frequent is M s ; only in the Cofan is it exceeded by M S . The Noanama are strikingly
frozen clot.
Electrophoretic conditions, buffers and different from the other three groups, with
substrate procedures for the enzyme and an N s frequency of 28.4% more than twice
serum protein tests were identical with that of the Cofan, and more than four
those described previously (Malcolm et al., times that of the Ingano: the Noanama
frequencies are similar to those for the
'72).
Gene frequencies were estimated using Cayapo in Brazil reported by Salzano et al.
the maximum likelihood program written ('72a). The tabulation of Post et al. ('68)
for us by W. J. Schull. Though the series and Salzano et al. ('72a) shows wide Auccontained persons of varying degrees of tuations in the gene frequencies of the
relationship all individuals were given MNSs system in other South American
equal weight in the gene frequency calcu- Amerindian populations, but only Layrisse
lations, and tests for Hardy-Weinberg equi- et al. ('63) have reported values for this
system in Colombia. Their results are withlibrium assumed random mating.
in the ranges found for the groups studied
RESULTS
here.
Rhesus
Blood groups
The five Rh antisera -C, -D, -E, -e and
A total of 168 Noanama, 78 Cofan, 51
Ingano and 64 Siona Indians were tested -c were employed as well as -Cw . No sam-
307
SERO-GENETIC STUDIES IN COLOMBIA
TABLE 1
Phenotype distribution for eight red cell antigen systems among some Amerindian
populations in Colombia
Noanama
Ingano
Cofan
Siona
Group
-
No.
Percent
No.
Percent
ABO
A
B
No.
Percent
No.
Percent
3
5.88
4.69
3.12
92.19
0
168
100.00
78
100.00
48
94.12
3
2
59
Tot a1
168
100.00
78
100.00
51
100.00
64
100.01
6
26
53
2
13
49
3.57
15.48
31.55
1.19
7.74
29.17
14
36
10
17.95
46.15
12.82
4
19
19
2
7.84
37.25
37.25
3.92
10
4
12.82
5.13
1.19
10.12
2
2
2.56
2.56
3.92
3.92
3.92
1.96
6.67
36.67
31.67
1.67
10.00
13.33
2
17
2
2
2
1
4
22
19
1
6
8
168
100.01
78
99.99
51
99.98
60
1
0.60
91
66
4
6
54.17
39.29
2.38
3.57
2
38
31
1
6
2.56
48.72
39.74
1.28
7.69
29
20
56.86
39.22
4
36
18
5
6.35
57.14
28.57
7.94
2
3.92
168
100.01
78
99.99
51
100.00
63
100.00
114
54
67.86
32.14
69
9
88.46
11.54
46
5
90.20
9.80
42
18
70.00
30.00
168
100.00
78
100.00
51
100.00
60
100.00
Lewis
Le(a )
Le(a - )
34
134
20.24
79.76
1
77
1.28
98.72
1
50
1.96
98.04
2
58
3.33
96.67
Tot a1
168
100.00
78
100.00
51
100.00
60
100.00
Duffy
Fy(a )
Fyfa - )
56
12
92.86
7.14
60
18
76.92
23.08
44
7
86.27
13.73
57
3
95.00
5.00
Total
68
100.00
78
100.00
51
100.00
60
100.00
68
100.00
78
100.00
45
6
88.24
11.76
55
91.67
5
8.33
MNSs
MS
MSs
Ms
MNS
MNSs
MNs
NS
NSs
Ns
Tota1
Rhesus
R z R z (CCDEE)
RzR, (CCDEe)
RIRl (CCDee)
R1R2(CcDEe)
Rlr (CcDee)
RzR2(ccDee)
Total
100.
P
p1+
p1Total
+
+
Kell
Kp(b + ), k
Kp(a + ) KpCb
Kp(a ) Kp(b
+
+ ), Kk
+ ), k
Tot a1
168
100.00
78
100.00
51
100.00
60
100.00
Diego
Di(a + )
Di(a -
168
100.00
28
50
35.90
64.10
16
35
31.37
68.63
36
25
59.02
40.98
Total
168
100.00
78
100.00
51
100.00
61
100.00
308
KIRK, McDERMID, BLAKE, GAJDUSEK. LEYSHON AND MAcLENNAN
TABLE 2
G e n e f r e q u e n c i e s f o r red cell a n t i g e n s y s t e m s
listed in table 1
Gene
ABO
A
B
0
Noanama
Cofan
Ingano
Siona
,0299
1.oooo
1.oooo
,9701
.0237
,0158
,9605
MNSs
MS
Ms
NS
Ns
,1499
.5465
,0197
.2838
.4661
,3929
,0211
,1199
.2843
.5 784
.0784
.0588
,2917
,5833
,0250
,1000
Rhesus
R,(CDE)
R,(CDe)
Rz(cDE)
R 0 (cDe)
,7550
,2337
,0113
,0129
,7050
.2756
,0065
,7647
,2353
.0335
.7839
.1411
,0415
P'
,4331
,6603
.6869
.4523
Lewis
Lea
,2024
,0128
.0196
.0333
Duffy
FYa
.7327
.5196
.6295
,7764
1.oooo
1.oooo
1.oooo
1.oooo
,9412
,9412
1 .oooo
KPh
Diego
Di a
0.0000
gene frequencies are higher than those reported by Layrisse et al. ('63) in Colombia
but close to the values given for other
South American groups by Post et al. ('68),
and for the Cayapo in Brazil by Salzano
et al. ('72a).
Lewis
Tests with anti-Lea revealed a major difference between the Noanama and the
other three linguistic groups. Twenty percent of the former were Le(a +) compared
with only 1 to 3% in the other groups.
With few exceptions, such as the Xavante
of Brazil (Gershowitz et al., '67) most populations of Indians in South America are
almost entirely Le(a-). Salzano et al.
('72a) report nearly 8 % of the Brazilian
Cayapo to be Le(a +).
P
Kell
k
DUffY
The Fy(a ) frequencies ranged from
76.9% for the Cofan to 95.0% for the
Siona. Similar ranges in frequency are
found for other Amerindian populations:
Post et al. ('68), and Layrisse et al. ('63)
give almost identical values in Colombia.
Salzano et al. ('72a) found 92% of the
Cayapo in Brazil to be Fy(a ).
+
,9583
+
Kell
Tests with four antisera -K, -k, -Kpa and
-Kpb revealed that nearly all persons were
ple reacted positively with the latter: in Kp(b + )k, the commonest phenotypic comaddition no cdelcde (rr) persons were de- bination in all parts of the world. However
tected and the gene frequencies were cal- five of the 60 Siona were Kp(a ) Kp(b )k
culated assuming absence of the r gene. and six of 51 Ingano were also K. This latAmong all four groups the RIRl and ter combination (Kp(a +)Kp(b +)Kk is unR I % phenotypes accounted for 85% or usual since the Kk, Kp(a)Kp(b) gene pairs
more of the total. The R 1 gene frequency appear to be inherited independently. The
fell within the range 0.71 to 0.78, with occurrence of six Kp(a ) persons as also
R2 in the range 0.14 to 0.28. R" is absent K in the Ingano can be explained only as
among the Noanama and Ingano, and only the result of the Kp" and K genes occurachieves a frequency of 0.01 in the Cofan, ring in the Cis conflguration in a group of
and 0.03 in the Siona. Like the MNSs sys- closely related individuals.
tem the rhesus system is characterized by
Diego
wide variations in frequency among Amerindian populations. Even in Colombia the
The Diego blood group system was first
populations studied by Layrisse et al. ('63) discovered among Amerindians in South
were more widely divergent than those re- America (Layrisse et al., '55) and it was
ported here.
proved to be an interesting marker system
among Amerindian populations, as well as
P
among Mongoloid populations in other
Tests with anti-PI revealed positive reac- parts of the world. In the present instance
tions ranging from 68% among the Noa- all persons were tested with Anti-Dia and
nama to 90% among the Ingano. The Pya some with the antithetical antibody -Dib.
,1994
,1716
,3598
+
+
+
SERO-GENETIC STUDIES IN COLOMBIA
Of those tested all were found to be Di
(b +). The frequency of Di(a +) persons
however was 0 % among the Noanama, 31
and 36% among the Ingano and Cofan
respectively and 59% among the Siona.
Similar ranges have been found in other
studies summarized by Post et al. (‘68).
Salzano et al. (’72a) report frequencies
close to 40% for Di(a ) persons in various
Cayapo groups in Brazil.
+
Serum protein and red cell enzyme
groups
The phenotype distributions for ten red
cell enzyme and four serum protein groups
among the Noanama, Cofan and Ingano
Indians are given in table 3. Some serum
samples were deficient in quantity so that
the total number of serum protein group
determinations was less than that for the
red cell enzymes groups. In addition one
red cell sample was exhausted before the
checking of PGM groups was completed.
Because of the conditions under which
the samples were collected and the elapsed
time before arrival in Canberra no attempt
was made to type for glucose-6-phosphate
dehydrogenase deficiency. All other systems, however, gave reproducible results
and we consider the likelihood of errors in
typing to be negligible.
Gene frequencies for those systems revealing variation, except for LDH which
will be discussed in more detail below, are
given in table 4. X’ values calculated on
the assumption of Hardy-Weinberg equilibrium revealed no significant departure
from expectation in any system.
Only four of the ten red cell enzyme systems examined revealed phenotypic variation in the three populations studied, and
of these only two were polymorphic by the
usual definition of having a variant gene
frequency of 0.01 or higher. In the red cell
acid phosphatase system three phenotypes,
A, AB and B were detected, and the P“
gene frequency did not differ significantly
among the three populations, having a
mean value of 0.22. This is comparable
with the value given by Salzano et al.
(‘72b) for the Cayapo (0.21) and by Tashian
et al. (’67) for the Xavante in Brazil (0.19)
but very much higher than the 0.02 among
the Yanomama (Arends et al., ’67) or the
0.05 for the Makiritare in Venezuela
(Weitkamp and Neel, ’70).
309
For locus-1 of the phosphoglucomutase
system three phenotypes 1-1, 2-1 and 2-2
were present and the P G M ’ , gene frequency of 0.78 again was almost identical with
that for the Cayapo in Brazil (0.76) given
by Salzano et al. (‘72b). Much higher values (0.94 and 0.85) were found respectively for the Yanomama and Makiritare in the
papers cited above, although in the present study the value of 0.84 for the Cofan is
not appreciably different than the value
for the Makiritare in Venezuela. In the
6 - phosphogluconate dehydrogenase system electrophoretic variants were detected
only among persons belonging to the Noanama linguistic group, four of 170 individuals having the “common variant” heterozygote phenotype (Carter et al., ’68). The
low frequency of the PGDC (listed as PGDB
by American authors) appears to be a characteristic of Amerindians in South America. Salzano et al. (‘72b) found no variants
and Weitkamp and Neel (‘70) found five
heterozygotes in one Makiritare village
but none in a further six villages and
Tashian et al. (’67) found no GPGD variants among 185 Xavante in Brazil. We
have not observed a n y significant diminution of GPGD activity comparable to that
postulated
for
“PGD-Makiritare” by
Weitkamp and Neel (‘70).
The only other red cell enzyme system
which revealed any variation was lactate
dehydrogenase. All 294 samples tested have
been listed in table 3 as normal. Two samples among the Ingano linguistic group,
however, showed the “atypical” pattern
which is illustrated in figure 4. In both
samples a strong single band occurs anodally to isozyme 3, and it appears to be
specific for lactate as substrate. Although
in the approximate position of LDH-X
found in mature male gonadal tissue and
sperm, comparison with human sperm
preparations showed that the x-band and
the “atypical” component were not of
identical mobility. One of the propositi was
a female aged 16 years and the other a 15
month old male. In the absence of full
family data for the propositi the interpretation of the “atypical” LDH patterns remains obscure.
All the other red cell enzyme systems,
PGM-locus 2, adenylate kinase, malate
dehydrogenase, peptidase A, peptidase B
and indophenol “oxidase” revealed no elec-
310
KIRK, McDERMID, BLAKE, GAJDUSEK, LEYSHON AND MAcLENNAN
TABLE 3
P h e n o t y p e distribution for red cell e n z y m e u n d s e r u m p r o t e i n groups trmoizg some
A m e r i n d i m g r o u p s i t 2 Colombia
Noanama
Cofan
A l l groups
Ingano
System
No.
Percent
No.
Percent
No.
9
58
102
5.33
34.32
60 36
2
32
40
2.70
43.24
54.05
Total
169
100.01
74
99.99
6-Phosphogluconate
dehydrogenase
6PGD A
AC
Total
166
4
170
97.65
2.35
100.00
74
100.00
-
-
-
74
100.00
50
100.00
Phosphoglucomutase
(locus 1)
P G M , 1-1
2-1
2-2
96
68
5
56.80
40.24
2.96
51
22
1
68.92
29.73
1.35
25
20
5
Total
169
100.00
74
100.00
Phosphoglucomutase
(locus 2)
PGMz 1-1
169
100.00
74
Lactate dehydrogenase
LDH
Normal
170
100.00
74
Malate dehydrogenase
MDH
Normal
170
100.00
Adenylate kinase
AK
1-1
170
Peptidase A
P e p A 1-1
1-1 weak
Total
Acid phosphatase
AcPh
A
AB
B
Percent
No.
-
-
14
36
28.00
72.00
11
104
178
3.75
35.49
60.75
50
100.00
293
99.99
50
100.00
290
4
294
98.64
1.36
100.00
50.00
40.00
10.00
172
110
11
58.70
37.54
3.75
50
100.00
293
99.99
100.00
50
100.00
293
100.00
100.00
50
100.00
294
100.00
74
100.00
50
100.00
2 94
100.00
100.00
74
100.00
50
100.00
294
100.00
126
44
74.12
25.88
52
22
70.27
29.73
37
13
74.00
26.00
215
79
73.13
26.87
I
-
Percent
170
100.00
74
100.00
50
00.00
294
100.00
Peptidase B
P e p B 1-1
170
100.00
74
100.00
50
00.00
294
100.00
Indophenol oxidase
Oxidase Normal
170
100.00
74
100.00
50
00.00
294
100.00
155
100.00
68
100.00
38
3
92.68
7.32
26 1
3
98.86
1.14
C aeruloplasmin
CP
B
A B
-
-
-
-
155
100.00
68
100.00
41
100.00
264
100.00
Hap toglobin
Hp
1-1
2-1
2-2
“0”
Total
27
79
48
1
155
17.42
50.97
30.97
0.65
100.01
42
21
5
61.76
30.88
7.35
-
-
68
99.99
22
8
6
5
41
53.66
19.51
14.63
12.20
100.00
91
108
59
6
264
34.47
40.91
22.35
2.27
100.00
Transferrin
Tf
C
155
100.00
68
100.00
41
100.00
264
100.00
Albumin
155
100.00
68
100.00
41
100.00
264
100.00
Total
I
Two of the samples gave “atypical” LDH patterns. These are discussed in the text.
SERO-GENETIC STUDIES IN COLOMBIA
TABLE 4
Gene frequencies f o r red cell e n z y m e and serum
protein systems showing varintion i n table 1
Alleles
All groups
Noanama
Cofan
Ingano
0.225
0.775
0.243
0.757
0.140
0.860
0.215
0.785
0.988
0.012
1.000
1.000
-
-
0.993
0.007
PGM
(locus 1)
PGM,'
PGM12
0.769
0.231
0.838
0.162
0.700
0.300
0.775
0.225
CP
C P
1.000
1.000
-
0.963
0.037
0.994
0.006
0.432
0.568
0.772
0.228
0.722
0.278
0.562
0.438
Ac.Ph.
Pa
Pb
6PGD
PGD A
PGDC
CPA
HP
HP'
H p'
-
trophoretic variants. Peptidase A showed
variation in intensity of the normal 1-1
band, some being classified subjectively as
weak. We have noted such variation repeatedly for samples collected under field
conditions, and although there may be a
genetic basis for part of the variation we
believe that much is due to instability of
the peptidase A itself.
Four serum protein systems, haptoglobin, transferrin, caeruloplasmin and albumin were examined: only haptoglobin was
polymorphic in the total population. The
H p ' gene frequency of 0.43 for the Noanama was significantly lower than for the
Cofan and Ingano linguistic groups (0.77
and 0.72 respectively). The overall Hp'
gene frequency of 0.56 is again almost
identical with the Cayapo frequency of
0.58 given by Salzano et al. ('72). The latter authors summarize the H p l gene frequencies for 73 series of Amerindians in
South America, giving a range of values
from 0.21 to 0.89, with all but two values
(Yupa and Guayakis) falling in the range
0.34-0.89. For transferrin the situation is
more complex. Nearly half the series of
Amerindian populations studied in South
America reveal the presence of genetic
variants other than Tf C. Some of these
variants are claimed to be Tf D1 similar to
that present in black American populations, others are designated Tf Dchi, which
311
is widely distributed in Mongoloid populations in other parts of the world. Kirk ('68)
has pointed out, however, that in many
cases no critical discrimination of the
transferrin variant found in a particular
population sample has been made so that
comparisons between Amerindian groups
on the basis of type of transferrin variant
present must be treated with caution. In
the present study only Tf C was detected.
Among South American Amerindians
genetic variation at the caeruloplasmin
locus has been investigated by Arends et
al. ('67, '70) and Salzano et al. ('72b). No
variants were reported for the Yanomama
and Makiritare in Venezuela, but Salzano
and his colleagues found 9 % BA heterozygotes among the Cayapo in Brazil. In our
own study we have found three persons, a
mother and two of her children, among the
Ingano linguistic group who were BA
heterozygotes. Careful comparison of these
phenotypes using the slab acrylamide technique of McCombs et al., ('70) as well as
using the standard starch gel technique
could not differentiate the A band from
that of a BA control (kindly supplied by
Dr. D. G. Shreffler). It is possible that our
own variant is identical with that found
among the Cayapo.
The other system, serum albumin, revealed no phenotypic variation. Albumin
variants have been found in three of the
four South American Amerindian series
investigated previously, only the Cayapo of
Brazil having no variants. Superficially
this suggests another similarity between
the present series and the Cayapo. However, the Albumin variant found among
the Warrau by Arends et al. ('69) and
among the Makiritare by Arends et al.
('70) is claimed by the latter authors not to
be detectable after starch-gel electrophoresis in the standard buffer systems used
for detecting albumin variants (Weitkamp
et al., '69). Since we have not employed
the buffer system used by Arends and his
group we cannot exclude the possibility
that variants similar to Albumin-Warrau
or Albumin-Makiritare are present in the
linguistic groups studied in the present investigation.
DISCUSSION
The detailed blood group, serum protein
and enzyme group studies reported here
312
KIRK, McDERMID, BLAKE, GAJDUSEK, LEYSHON AND MAcLENNAN
LDH 1
LDH 2
+
t
LDH 3
c.
LDH 4
Origin
a.
b.
c.
d.
Fig. 4 “Atypical” LDH patterns from two persons belonging to the Ingano linguistic group
(b and c) compared with normal controls (a and d).
for Amerindian populations in Colombia
are of interest from several points of view.
Firstly, they may be used to indicate the
degree of genetic similarity between the
populations sampled in the two broad regions studied, the Noanama (including the
very small sample of Empera) from the
Pacific lowlands, and the three groups
from the Putumayo, each of which in turn
is relatively isolated from the others. Secondly, the results may be used to compare
the Colombian populations with those in
other parts of South America for which
comparable data is available. Finally, studies of this kind are of value for the information they contribute on the amount of
genetic heterogeneity present in small
human populations, including the occur-
SERO-GENETIC STUDIES I N COLOMBIA
rence of specfic mutants with localized
distributions.
With respect to the first of these points,
our data indicates the distinctiveness of
the Chocoan-speaking Noanama from the
Putumayo groups, and this is demonstrated
in a number of systems. For example, the
Noanama have only 15% M S but 28% N s
in the MNSs system in contrast to the
range of M S from 28 to 47% and N s from
6 to 12% for the Putumayo groups. Similarly the Lc" frequency for the Putumayo
groups has a range from 1 to 3 % , but is
20% in the Noanama and the latter have
only 43% for the Hpl gene whilst the two
Putumayo groups studied have 72 and
7 7 % . The Noanamo also do not have the
Di" gene, whereas the Di" frequency
among the Putumayo groups is 17 to 36 % .
There are some marked differences between the Putumayo groups themselves.
This is not surprising perhaps, since the
three groups studied for the red cell antigen systems (two of which were investigated also for the cell enzyme and serum
protein systems) provide little evidence for
intermarriage between them and belong to
distinctive linguistic families. However,
there is no consistent pattern for the differences between the three Putumayo
groups so that on the basis of the present
evidence we cannot pick out the Quechuaspeaking Ingano for instance a s being quite
different from the Cofan and Siona. Also,
since data is not available for Quechuaspeakers from the Colombian Altiplano
we can add nothing of value toward solving the problem of the relationship between
the highland and lowland Quechua speakers
One further point of interest is the possible influence of non-Amerindian populations on the gene pool of the Putumayo
groups. As emphasized in the section on
the ethnography of these groups they are
surrounded by numerically superior Mestizo and Black populations. With the exception of the ABO and possibly the Rhesus
blood group systems the evidence indicates
little if any introduction of alien genes into
either the Pacific or Putumayan populations studied here. In the ABO system the
Ingano have 3% of A genes whilst the
Siona have nearly 4% of A and B genes
combined. The Noanama and Cofan have
only 0 genes, a situation considered to be
characteristic of Amerindian populations
313
in South America in the pre-contact period. In the Rhesus system the R gene
complex present in the Cofan and Siona
groups may be also introduced from alien
sources. By contrast, a number of genes
characteristic of European or Black African populations were not detected in any
of the Pacific or Putumayan groups. These
genes include the r(cde) gene of the Rhesus system, U negative in the MNS system,
P C and PR in the red cell acid phosphatase
system, variants at the PGM second locus
and slow-moving DI variants in the transferrin system. The absence of such specific
variants does not rule out the possibility
of intermarriage between the Amerindians
and their alien neighbours, but suggests
either that such intermarriage is infrequent or that the hybrid offspring are not
identified with the parent Amerindian
group. The ethnographic evidence supports the latter suggestion, at least for
some of the Putumayo groups.
If we turn now to an examination of the
genetic relationship between the Colombian groups studied here and Amerindian
populations in other parts of South America
we are faced by the paucity of information
for a wide range of genetic markers in
order to make a detailed comparison possible. The high frequency of the blood
group 0 gene in the ABO system and the
very variable frequencies between the different linguistic groups for many of the
genes in some of the other blood group
systems such as Ns, R. and Di" are comparable to the situation reported for Amerindians in other parts of South America.
What is of interest is that for the serum
protein and red cell enzyme systems which
are polymorphic the frequencies of the
genes are very similar to those reported for
the Cayapo in Brazil, but different in several respects from the results found by
other investigators for Xavante in Brazil or
the Yanomama and Makiritare in Venezuela as noted in the preceding section. The
blood group results also are similar in a
number of respects to these reported for
the Cayapo of Brazil. When more information is available the significance of these
relationships may become clearer: we intend to publish a more detailed analysis
in the future.
Finally we wish to note some points of
interest about the results of our present
314
KIRK, McDERMID, BLAKE, GAJDUSEK, LEYSHON AND MAcLENNAN
analysis with respect to the genetic heterogeneity of small human populations, and
the occurrence of mutations with restricted local distribution. The genetic heterogeneity for the systems studied here is low
in contrast to that reported for European
and Black African populations (Harris and
Hopkinson, ’72). This is true for the ABO,
Rhesus and Kell blood group systems, and
for the serum protein and red cell enzyme
systems. Only two of the ten red cell enzyme systems and one of the four serum
protein systems are polymorphic by usual
standards in all the three linguistic groups
for which information is available, though
a n additional red cell enzyme system is
polymorphic among the Noanama and an
additional serum protein system is polymorphic among the Ingano. This lack of
heterogeneity is characteristic of peripheral non-human populations and unpublished information from one of our laboratories indicates that this is true also for
peripheral human populations. The Pacific
and Putumayo groups therefore may be
considered as small genetic isolates with
increased genetic homogeneity, a condition
which may be further increased if their
culture does not permit the retention of
genetic variability introduced from sources
outside the group. In this connection it is
interesting that we detected no new mutants in a n y of the systems investigated,
with the exception of the two persons with
an “atypical” component in the lactate
dehydrogenase system. This unusual LDH
variant merits more detailed comment.
Comparison with the LDH-X present in
sperm from Caucasian controls showed
the “atypical” component to have a slightly different mobility, but it would be of
interest to compare directly with the LDHX present in extracts of sperm from males
belonging to the Cofan linguistic group. It
seems unlikely, however, that the LDH-X
of Amerindians in Colombia would be different in its electrophoretic behaviour from
that present in Caucasians. Moreover, if
the “atypical” band and the Cofan LDH-X
were identical it would be necessary to postulate that the gonadal isozyme was derepressed in the red cells of the two persons
encountered in the present study. It is more
tempting to postulate that the “atypical”
component in these two persons is related
to some disease process, but further study
is needed to establish if this is correct. It is
relevant to note that in nearly 10,000 red
cell samples examined for electrophoretic
variants in one of our laboratories during
the last few years only one analogous
“atypical” pattern has been detected. This
was found in a n apparently healthy Chinese in Singapore, where a single “atypical” component was present in the LDH
pattern migrating slightly slower than isozyme 2 (Blake et al., ’73).
ACKNOWLEDGMENTS
We are indebted to Mrs. Elizabeth Robertson, Mrs. Kathy Sims and Mr. Robert
Taylor for excellent clerical and technical
assistance during the conduct of this research. We also thank Dr. Nicholas Escobar, Director of the Texaco Company in
Colombia, and his staff at Orito for helicopter transport and other logistics support which he made available to us in the
Putumayo region. We are grateful to Sr.
Ugo Cuevas of Cali, Colombia for sea transport from Buenaventura to Belem on the
Rio Docampado and the return voyage in
one of his shrimping vessels.
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