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Blood groups red cell enzymes and cerumen types of the Ahousat (Nootka) Indians.

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Blood Groups, Red Cell Enzymes, and Cerumen
Types of the Ahousat (Nootka) Indians
BRAXTON M. ALFRED, T. D. STOUT, JOHN BIRKBECK, MELVIN LEE
AND NICHOLAS L. PETRAKIS
Department of Anthropology and Sociology, University of British Columbia;
Canadian Red Cross, Vancouver, B. C.; School of Home Economics,
University of British Columbia; Department of Epidemiology and
International Health, University of California,
San Francisco, California
A survey of the blood groups of a Nootka band produced frequencies
ABSTRACT
characteristic of North American Indians for the ABO system (0.99 for 0, 0.0 for B,
and 0.01 for A), Rhesus (0.822 for cDE, 0.011 for cde, 0.023 for cDe), Lutheran
(1.00 for Lu(a-)), Duffy (0.505 for F y f a f ) ) and Diego (0.039 for Di(a+)). K is
not absent though the frequency is not great (0.028). Surprising results were obtained for the MN locus (0.399 for M, 0.601 for N ) , P (0.209 for PI), and Lewis
(0.568 for Le(o+)). A frequency of phosphoglucomutase type P G M I ~of 0.890 was
found; all hemoglobins were type AA; n o G-6-PD deficiency was found a n all were
type B positive; the frequency of the dry cerumen allele was found to be 0.323.
The Ahousat are a Central Nootkan
group currently living on Flores Island
which is off the west coast of Vancouver
Island somewhat north of Barclay Sound
(fig. 1). Earlier their primary concentration was on the mainland near what is now
called Chetarpe with a subsidiary location
at Ahous Point on Vargas Island. They
came by their present location after a long
and bloody war with the Otsosat early in
the nineteenth Century.
In 1774 a Spanish vessel put into Nootka
Sound; Cook arrived in 1778 and his visit
was responsible for beinning the lucrative
sea otter fur trade with China. This contact brought to the Nootka iron, copper,
brass, muskets, cloth, rum and disease.
Small pox epidemics ravaged the population periodically with a particularly severe outbreak in 1852. Also measles, influenza, tuberculosis, venereal disease and
alcohol were newly available means of
death. Aboriginally the west coast of Vancouver Island supported one of the greatest population densities in the New World;
the impact of contagious diseases was
magnified correspondingly. The social
structure was greatly affected because “Decrease of population meant that chiefs
died without heirs in the direct line.”
(Drucker, ’51: 12).
AM. J.
PHYS.
ANTHROP.,31: 391-398.
In the 1850’s the dogfish oil trade arose
with the nascent lumbering industry. In
1860 the town of Alberni was established
at the head of the Alberni Canal in the
region of the Sproat Lake Nootkan groups;
in 1875 Father Brabant’s mission was
established at Hesquiat. The fur seal trade
blossomed in the 1880’s and some Nootka
men, shipping as hunters, made visits to
Japan, China and San Francisco. Canneries began to appear and the town of Tofino
was established around the turn of the
century.2
Reasonably good population counts
have been made since about 1880. Figure
2 presents the population curve for the
Ahousat band since 1883. The curve drawn
in is not fitted; it is to be used only as a
guide to the likely underlying trend. Between 1965 and 1968 the population of
registered Indians at Ahousat increased by
8.05% or roughly 4% each year. The
shape of the population curve at Ahousat
is very similar to that reported by Duff
(’65: 44) for the total provincial Indian
population.
We are, then, considering a group of
people having survived 200 years of white
1 We wish to thank Dr. Bruce Chown, Rh Laboratory
Winnipeg, Manitoba for reading and commenting on’
an earlier draft.
2 This account is drawn heavily from Drucker (’51)
and Duff (’64).
391
392
ALFRED, STOUT, BIRK3ECK, LEE AND PETRAKIS
\\
COURTENA
cL*yoQfio~
C
O
<
Figure 1
.-
Kelienat
lJdgOmOied
Z
100
1880
'
1890
1900
1910
1920
1930
1940
' L
1950
1960
1' '0
Y E A R
Figure. 2
Population trend at Ahousat
BLOOD FACTORS AND CERUMEN TYPES OF
contact. Northwest coast warfare often
involved taking women and children as
slaves. This, plus the pressure of band
exogamy should have resulted in genetic
mixing sufficient to obscure any local effects of white contact; that is local effects have either diffused through the region or have been cancelled.
In March 1968 a survey of nutrition
and health was conducted at Ahousat under the direction of Dr. Melvin Lee, University of British Columbia School of Home
Economics. Blood was drawn routinely and
typing carried out by the Candian Red
Cross. The blood was also studied for
hemoglobin, glucose-6-phosphate dehydrogenase (G-6-PD) and phosphoglucomutase
(PGM) polymorphisms. In addition the
subjects were classified as to cerumen
type, a recently described genetic polymorphism.
METHODS
A. Sampling
Subjects were not randomly drawn and
no attempt was made to exclude relatives.
Of the 669 registered Indians on the
Ahousat band list, 310 were living on the
Reserve at the time of the survey, Blood
samples were obtained on 198, 64% of
those in residence. Pedigrees3 were constructed; from these we estimate that three
of our subjects are not related closely
(i.e. as sibling or as parent/child) to another member of the sample. (No first
cousin marriages appear.) The amount of
relatedness in the sample is not serious
concern since this characterizes the (finite)
population for which genetic parameters
are to be estimated.
The sampling technique, however, may
be a serious deficiency although intuitively
it does not seem that it should be. In
random sampling from a finite population
without replacement, at the kth draw there
are N - k 1 items remaining; therefore,
each must have probability 1/N - k 1
of being included. When subjects are selfselected, which was true of this survey
and probably of nearly all such surveys,
this probability does not obtain. For the
present we will make the assumption that
our sample is the equivalent of a random
sample.
+
393
THE NOOTKA
From figure 3, which presents the age
by sex distributions for the Ahousat band
list, and the current sample, one may see
that the sample is fairly representative of
the population with regard to these two
attributes.
B. LaboTatoy
All blood specimens were collected in
heparin. Most of the serum was removed
from each specimen for biochemical studies and ACD was then added to the cells.
This was done within 24 hours of collection. The ACD specimens were then refrigerated and shipped to the Red Cross
Blood Transfusion Senice in Vancouver.
The cells were grouped during the next
four days4 There was no appreciable hemolgsis during the test period.
The anti-sera and grouping techniques
are given in table 1.
A number of cells were retested as
follows:
Nine e-negative specimens were checked
with two anti-e sera by three methods and
Y t R
FEMALE
&*NO
,
SAMPLFI-J
+
Fig. 3
Age, sex population tree.
SWe wish to express our gratitude to Mr. George
Pettifer Public Health Nurse, Medical Services, of
Tofino $or providing much of the information necessary to complete and correct pedigrees.
4Typing was carried out by Mrs. Isabella Horvath,
R.T. and Mrs. Judith Decker, R.T. of the Red Cross
Elood Transfusion Service, Vancouver, B.C.
394
ALFRED, STOUT, BIRKBECK, LEE AND PETRAKIS
TABLE 1
Anti-sera and grouping techniques
Anti-sera
Anti-A1
Anti-A
Anti-B
Anti-A & B
Anti-D
Anti-C
Anti-E
Anti-?
Antie'
Anti-M
Anti-N
Anti-S
Anti2
Anti-P
Anti-Lua
An ti-Lub
An ti-Lea
Anti-Leb
Anti-Kell
Anti-k
Anti-Fya
Anti-Fyb
AntiJKa
Anti-JKb
Anti-WP
Anti-Wr"
Anti-Di"
Anti-Vel
An ti-Vel
Anti-Human Globulin
30% Bovine Alb.
Source
Sal.
Sal.
Sal.
Sal.
Sal.
Sal.
Sal.
Inc.
Sal.
Sal.
Sal.
Can. Red Cross B.T.S. Toronto
Can. Red Cross B.T.S. Toronto
Can. Red Cross B.T.S. Toronto
Can. Red Cross B.T.S. Toronto
Can. Red Cross B.T.S. Toronto
Can. Red Cross B.T.S. Toronto
Can. Red Cross B.T.S. Toronto
Can. Red Cross B.T.S. Toronto
Ortho Pharmaceutical Corporation
Ortho Pharmaceutical Corporation
Can. Red Cross B.T.S. Toronto
Can. Red Cross B.T.S. Toronto
Blood Grouping Labs., Boston
Can. Red Cross B.T.S. Toronto
Can. Red Cross B.T.S. Toronto
Dr. M. N. Crawford
Can. Red Cross B.T.S. Vancouver
Can. Red Cross B.T.S. Vancouver
Can. Red Cross B.T.S. Vancouver
Can. Red Cross B.T.S. Vancouver
Can. Red Cross B.T.S. Vancouver
Dade (Dried)
Can. Red Cross B.T.S. Vancouver
Can. Red Cross B.T.S. Toronto
Can. Red Cross B.T.S. Toronto
Can. Red Cross B.T.S. Vancouver
Dr. Bruce Chown, Winnipeg
Can. Red Cross B.T.S. Toronto
Can. Red Cross B.T.S. Vancouver
Pfizer
Pentex
Test
Capillary
Capillary
Capillary
Capillary
Capillary
Capillary
Capillary
Lows Pap. Capillary
Lows Pap. Capillary
Lows Pap. Capillary
Lows Pap. Capillary
Ind. Coombs. Cap.
Ind. Coombs. Cap.
Capillary
Capillary Lows Pap.
Ind. Coombs
Capillary
Saline Tube
Capillary Lows Pap.
Capillary Lows Pap.
Ind. Coombs Cap.
Ind. Coombs Cap.
Ind. Coombs Cap.
Sal. Ind. Coombs. Cap.
Ind. Coombs Cap.
Method for
re-check neg.
results
I.C. Tube
I.C. Tube
Sal. Tube
Sal. Tube
Sal. Tube
I.C.
I.C.
I.C.
I.C.
Tube
Tube
Tube
Tube
Ind. Coombs. Cap.
Ind. Coombs. Cap.
Capillary
Capillary
I.C. Tube
Tube
One hundred fifty-four blood samples
including two positive controls, one of
which was an Ahousate positive specimen. were analyzed electrophoretically for heThis was done to establish that the anti-e moglobin, G-6-PD, and phosphoglucomutase types employing methods described by
was working properly.
Because of the unusual MN findings Briere et al. ( ' 6 5 ) , Kirkman and Hendrickthirty secimens were sent to the National son ( ' 6 3 ) , and Spencer et al. ('64), reLaboratory, Canandian Red Cross Blood spectively.
Cerumen type was determined by visual
Transfusion Service, Toronto. The results
in the National Laboratory were identical inspection employing an otoscope accordwith those in Vancouver. The National ing to the criteria given by Petrakis ('67).
Laboratory also tested 5 M M and 5 N N
RESULTS
cells of this group with anti-Mc, anti-Mur
In
table
2
are
presented allele and pheand anti-Mi". All 10 cells were negative
notype frequencies. There are no large
with these three antisera.
All P negative cells were retested with departures from expectation. Consequently,
anti-PI to check as to whether any weakly it appears that the mating pattern of the
reacting P cells had been missed. There population results in the stochastically independent union of genes at all loci obwere no discrepancies.
The three group A individuals were served.
In table 3 are presented the Rh and
tested with anti-Al and were all AI.
The six (Le(a-,b-)
cells were re- MNSs chromosone frequencies; the former
checked using saline tube test and found were computed by an iterative procedure
which will be described elsewhere.
to be correct.
BLOOD FACTORS AND C E R U M E N TYPES O F
395
THE NOOTKA
TABLE 2
Blood group phenotype and gene frequencies
System
ABO
P
Phenotype
A
B
AB
0
Pi
Pz
Kell
Diego 2
Duffy a
Kidd
Lewis 5
MNSs
Rhesus
kk
Kk
KK
Di( a + )
Di(a-)
FY(a+ )
Ma-)
Jk( a+ 1
Jk( a- 1
W a f )
Le(a-)
MMSS
MMSs
MMss
MNSS
MNSs
MNss
NNSS
NNSs
NNss
CcDEe
CcDEE
ccDEe
ccDEE
ccDee
CcDee
Observed
phenotype
frequency
Allele
3 (0.015)
o (0.000 j
0 (0.000)
195 (0.985)
74 (0.374)
124 (0.626)
187 (0.944)
11 (0.056)
0 (0.000)
12 (0.075)
148 (0.925)
184 (0.929)
14 (0.071)
118 (0.596)
80 (0.404)
113 (0.571)
85 (0.429)
0 (0.000)
l(0.005)
24 (0.121)
l(0.005)
20 (0.101)
87 (0.439)
0 (0.000)
8 (0.040)
57 (0.288)
27 (0.136)
19 (0.096)
8 (0.040)
140 (0.707)
l(0.005)
3 (0.015)
Allele
frequency
A
B
0
0.008
0.000
P1
PZ
k
K
0.209
0.791
0.972
0.028
D1(
0.992
a+)
Di(a-)
FY(a+ 1
Ma-)
Jk(a+)
Jk(a-)
W a f )
Le(a-)
M
N
S
S
D
C
E
C
e
0.038
0.962
0.734
0.266
0.364
0.636
0.756
0.244
0.399
0.601
0.078
0.922
1 .ooo
0.124
0.891
0.876
0.109
All (3) A positive reactions were to anti-AI, i.e. no A2 was found.
No further testing.
aAnti-Fy(b) was used on 30 bloods; 10 were Fy(a+,b+), 16 were Fy(a+,b-1,
and 4 were
Fy(a-.b+).
*Anti-Jk(b) was used on 30 bloods; 17 were Jk(a+,b+), 1 was Jk(a+,L+).
and 12 were
Jkf a--.b4->.
- 5 A l l subjects were more than 18 months old, Anti-Le(b) was also used; 113 were Le(a+,b-),
79 were Le(a-,b+), and 6 were Le(a- b-).
Additionally 92 were typed for WrigGt and all were Wr--; 195 (all type 0 bloods) were typed for
Lutheran and all were Lu(a-,b+);
189 were typed for Vel and all were Vel+.
2
Hemoglobin, G-6-PD, electrophoresis,
and cerumen types.'. All hemoglobins were
type AA. All samples were type B positive
for G-6-PD and no deficiency was found.
Phosphoglucomutase (PGM) electrophoresis revealed 79.3% type 1-1 and 20.7%
type 2-1 giving allele frequencies f o r
PGM: of 0.89 PGMf of 0.11. Eighty-nine
of 244 subjects (32.3%) had dry type
cerumen giving q = 0.56.
The sample was broken into four age
groups to determine if there is any obvious
trend through time for the frequency o f
positive reactions to any of the anti-sera
used. The result is presented in table 4.
Since all the positive reactions to anti-A
are given by subjects under 20 years old
(i.e., born after 1948) we may tentatively
conclude either that the A antigen is a
recent introduction or that selective pressure on this locus has not had time to
eradicate it. Intuitively, the former seems
more likely as there is no presently known
relationship between this antigen and any
5 We gratefully acknowledge the assistance given
by Mr. Robert Mossberger and Dr. Luan Eng Lie-Injo
in the electrophoretic studies. This part of the work
was partially supported by a gift from Viola K.
Schroeder i n memory of Walter Schroeder.
396
ALFRED, STOUT, BIRKBECK, L E E AND PETRAKIS
TABLE 3
Chromosome frequencies
0.014
0.047
0.023
0.822
0.011
0.003
0.020
0.060
cDE (Rx)
Rhesus
cde (r)
CdE (R,.)
cdE ( R ’ )
MS
0.030
0.369
0.049
0.552
MS
MNSs
WS
NS
selective agent. As mentioned above, however, all A positive reactions were to type
AX and so this is not evidence for white
admixture. Except for P and Diego, most
proportions seem to be fluctuating randomly. At €’ we note that the proportion
with the PI antigen in the total sample
is 0.209; among those individuals born before 1908 it is 0.042 (i.e., the proportion
o f the PI phenotype is 0.083). Again, however, we cannot know whether there was a
large influx of the PI antigen after 1908 or
whether some unknown selective pressure
is operating. The latter seems less likely
because the proportions have apparently
stabilized. At Diego a similar kind of conclusion is reached; the proportion with the
DIa antigen is 0.03 for those born after
1928 and is 0 for those born before. We
do not wish push this discussion of pos-
sibilities beyond simply calling attention
to them as the numbers on which the proportions are based are too small to warrant
anything else (Gajdusek, ’64).
DISCUSSION
Two previous studies have reported on
Nootkan samples not mixed in with other
groups. The amount of directly comparable
information is small, however.
Hulse (’55: 96) reports gene frequencies of (0.02, 0, 0.98) for A, B and 0
respectively for his “Central Nootka” samplc6 Ride (Mourant et al., ’58: 190) got
(0.03, 0, 0.97) for the Nootka in 1935.
A, is absent in our sample and neither of
the two prior studies made the discrimination. At the Rh locus D, neither Hulse nor
6Professor Hulse h a s kindly suppljed us with his
original data. Among those he .fyped i n 1955, 14 were
carried on the Ahousat band list in Maxch 1968. Of
these, 4 appeared jn our sample. There was complete
agreement on the ABO typing, but discrepancies arose
at the MN and R h loci. First consider the results
of Mn:
Subject
Hulse
Ahousat
1
2
3
4
M
M
N
MN
M
N
MN
MN
In his 1955 paper.Hulse reports only the Rh locus D.
In the data he provided to us, however, information is
also given on C and E; presumably 3 anti-sera were
used. The following comparison is possible:
Subjcct
Hulse
Ahousat
I
2
3
cDE
CDE
CDE
CDE
CcDE
cDE
CcDEe
CcDEe
4
There i s complete agreement at the D and E loci
but discrcpancies at the C locus.
TABLE 4
Positive reactions to selected anti-sera by age group
Anti-serum
3
Total
sample
Less t h a n
20 years
N = 120
Between
20 and 40
years
N = 37
0.015
0.242
0.985
0.197
0.672
0.879
0.152
0.995
0.374
0.571
0.056
0.929
0.596
0.061
0.025
0.233
0.975
0.192
0.633
0.875
0.150
0.000
0.270
1,000
0.216
0.649
N=1871
1.000
0.417
0.600
0.058
0.950
0.642
0.058
This is the total on whom age was available.
0.865
0.162
1.000
0.297
0.459
0.054
0.892
0.541
0.081
Between
40 and 60
More t h a n
60 years
N = 12
0.000
0.222
1.000
0.111
0.833
0.889
0.167
1.000
0.444
0.611
0.056
1.000
0.500
0.000
0.000
0.167
years
N = 38
1.000
0.167
0.750
0.750
0.167
0.917
0.083
0.667
0.083
0.833
0.750
0.000
BLOOD FACTORS AND CERUMEN TYPES OF THE NOOTKA
the present study observed any cases of the
genotype dd.
Neel and Salzano ('64: 87) listed "Some
Distinctive Genetic Features of the American Indian" as follows:
System
High frequency of 0
High frequency of M
Rh
The frequency of cDE is higher than
any where else in the world; cde and
cDe low or absent.
Kcll
K may have been absent in
pre-Columbian Indians
Duffy
High frequency of F7a
Diego
High frequency of D,a
Lewis
Almost all populations show Le(a-)
only
Luthernn Almost all populations show Lu(a-)
ABO
MN
only
(Some notational changes have been introduced.)
To this list might be added that North
American Indians have very high frequencies of the allele for dry type cerumen
(Petrakis et al., '67; Petrakis, '69).
The blood group allele frequenices at
Ahousat are ambiguous. On the one hand,
the frequencies of the 0 and A, antigens,
the Rh pattern, and the uniformly Lu( a-)
traits are aboriginal Indian indicating little,
ii any, white admixture. But the MNSs
system frequencies stand in direct contradiction to, not only the Neel and Salzano
criteria, but also to every other study of
the Northwest Coast known to us. Further, the pattern bears little resemblance
to the English or Candian pattern as is
seen below.
English
1
Canadian 1
Ahousat
1
MS
0.247
0.255
0.030
Ms
0.283
0.304
0.369
NS
0.080
0.061
0.049
Ns
0.390
0.380
0.552
(Race and Sanger, '68: 91)
Chown and Lewis (Montagu, '60: 347)
report the following phenotypic proportions
for "British Columbia Indians :"
M N S S Phenotgpic Frequencies
Chown
Lewisand
MMSS, n'6MSs
MMss
MNSS, MNSs
MNSS
NNSS. NNS5
NNss
0,227
0.370
0.090
0.257
0.010
0 047
Ahousat
0.005
0.121
0.106
0.439
0.040
0.288
~-
Chown and Lewis used anti-sera M, N, S while the
Ahousat study used M, N. S and s.
397
At the locus P the frequency of the Pi
phenotype is 0.374, giving a gene frequency of 0.209. Montagu ('60. 356) reports phenotypic frequencies this low or
lower in Australia (0.23), Thailand (0.30)
and New Britain (0.35); Caucasian frequencies are around 0.78 for the Pi antigen
(Race and Sanger, '68: 159). Doeblin
and IClohn ('67: 706) report a frequency
for the Pi phenotype of 0.896 among the
Seneca, and Thomas et al. ('64: 192) report 0.637 and 0.582 at Masset and Skidegate respectively in the Queen Charlotte
Islands.
The Caucasian frequency for the Kf
phenotype seems to lie between about 0.05
to 0.09 (Race and Sanger, '68 : 266; Chown
et al., '63: 379). This may be compared
with 0.056 at Ahousat, which is intermediate among the range of studies reported by
Montagu ('60: 357) but considerably
higher than most other North American
Indians (e.g. Thomas et al., '64; Doeblin
and Mohn, '67).
Comparing the Diego locus with the
Indian samples reported by Laryisse ('58:
177, 182) it is seen that the Ahousat sample is clearly more typical of North than
of South American Indians in general. But
the Haida have a frequency for the D,"
antigen of 0.007 (Thomas et al., '65: 192),
and the geographically nearest comparable
frequency occurs among the Blackfoot of
Alberta (Doeblin and Mohn, '67: 706), an
Algonkian speaking group; next are the
Southwest Athapaskans, i.e., Apache 0.041,
Navaho 0.047. Similarities with the Apache
end, however, with this Diego frequency
(Gershowitz, '51 : 197). Apparently there
are no similar frequencies to the north,
e.g. Alaskan Athapaskan, 0.005, Tlingit,
0.000, Kutchin, 0.000 (Doeblin and Mohn,
'67).
Finally, a very striking frequency was
obtained at Lewis. The Ahousat frequency
for the Le ( a + ) phenotype is 0.571. With
the exception of a Central Australian
group, which was 100% Le(a+), reported
by Simmons et al. (Montagu, '60: 361)
this is the highest frequency known to us.
Thomas et al. ('64: 192) found 0.239
and 0.232 at their two sites, Doeblin and
Mohn ('67: 706) found 0.208 among the
Seneca. The only region that seems to have
398
ALFRED, STOUT,
BIRKRECK, LEE
AND PETKAKIS
Negroes; Blood Types of Asiatic Indians and
frequencies of this phenotype in the range
Apaches. Am. J. Phys. Anthrop. 17 (3): 195observed at Ahousat is Polynesia (Montagu,
200.
'60: 361).
Giblett, E. R. 1967 Variant Phenotypes: HaptoAnalysis of phosphoglucomutase (PGM)
gobin, Transferrin, and Red Cell Enzymes. In:
Advances in Immunogenetics. T. J. Greenwalt,
types demonstrated a high frequency of the
ed. J. B. Lippincott.
PGM: (0.89), which is similar to that
found in Athapaskan and Aleut Indians of Hulse, F. S. 1955 Blood-types and Mating Patterns Among Northwest Coast Indians. SouthAlaska, Scott et al. ('66), and in the Ainu
west J. Anthrop., 11: 93-104.
in Hokkaido, Giblett ('67), and distinctly Kirkman, N. H., and E. Af. Hendrickson 1963
higher than the frequencies reported for
Scx-linked electrophoretic difference in glucoseF-phosphate dehydrogenase. Am. J. Hum. Gen.,
Orientals, Caucasions and Negroes,
15: 241-258.
Spencer et al. ('64). It has been previously
M. 1958 Anthropological considerasuggested by one of us (N.L.P.) that in- Layrissc,
tions of the Diego (Dia) Antigen. Am. J. Phys.
creased frequency of the wet (sticky)
Anthrop., 16 ( 1 ) : 173-186.
type of cerumen present in Indian tribes Matsunaga, E. 1962 The dimorphism i n human normal cerumen. Annals of Human Gemight be indicative of admixture with
netics, 25: 273-286.
Caucasians, since Indian tribes known hisM. F. A. 1960 Introduction to Phystorically to be unmixed have higher fre- Montagu,
ical Anthropology. Charles C Thomas, Springquencies of the dry allele than found in
field, Illinois,
this study. Our present findings on blood Mourant, A. E., A. C. Kopec and K. DomaniewskaSobczak 1958 The ABO Blood Groups. Blackgroups and PGM types of the Ahousat do
well Scientific Publications, Oxford.
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