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Blood groups and haptoglobins in the Eastern Carolines.

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Blood Groups and Haptoglobins in the Eastern Carolines 1
NEWTON E. MORTON AND MANABU YAMAMOTO
Population Genetics Laborutory, University of Hawaii, 241 1 Dole Street,
Honolulu, Hawaii 96822
KEY WORDS
Blood groups . Haptoglobins
. Micronesia.
ABSTRACT
The ABO, M N S , Rh, and haptoglobin systems were examined in
samples from the populations of Pingelap, Mokil, Ponape, and Kusaie. There is
marked differentiation between neighboring islands.
Although there has been extensive blood
grouping in Micronesia, the Eastern Carolines have been little studied. As part of
a n investigation of population structure
we collected bloods from natives of Pingelap and Mokil atolls and the volcanic
islands of Ponape and Kusaie. The systems ABO, MNS, Rh, and haptoglobins
were examined. Studies on isozymes and
immunoglobins are reported elsewhere
(Yamamoto and Fu, '72; Steinberg and
Morton, '72).
MATERIALS A N D METHODS
There is a good airline service between
Ponape and Honolulu, but the other
islands of the Ponape District are reached
by a ship which anchors just long enough
to leave supplies and load copra and
handicrafts. Since refrigeration on the
atolls is poor, we had to limit venepuncture to visits by ship, when many islanders are busy. We collected as many bloods
as we could during the two field trips,
but it was impossible to obtain a n exhaustive or even random sample by venepuncture. Cells for typing were collected
in ACD vacutainers (referred to as the
lab sample in tables 1-3) and kept refrigerated until receipt in Honolulu, up
to three weeks later (Yamamoto and Fu,
'72). After most of the cells had been
glycerolized, each tube was half-filled with
Alsever's solution containing 0.1 % sodium azide. During the following week
tests were run i n duplicate, by standard
saline and ficin methods (Morton, '64)
and a Microtiter technique modified from
Wegmann and Smithies ('66). Inconsistent results were repeated. Saline and
enzyme-potentiated reactions were strong,
A m J. PHYS. ANTHROP.,38: 695-698.
but cell fragility made the repeated washings for a Coombs' test impractical. Although there was good recovery of cells
from glycerol by dialysis against citrate
(Morton and Rosenfield, '67), these frozen
cells were rarely required except for isozyme typing, where separation from glycerol is unnecessary.
During the first field trip in 1969 we
experimented with typing of capillary samples drawn from nearly all Pingelapese
and Mokilese on the atolls, and a substantial fraction of migrants to Ponape
(referred to as the field sample in tables 1-3). A small drop of blood was
drawn from the finger by a disposable
lancet into 2 ml of 0.85% saline, giving
a faintly pink suspension. Commercial
antisera were diluted in a solution containing 25 mg polyvinylpyrolidone K-90
(PVP), 10 ml of 22% bovine albumin,
one drop Tween 80, 989 ml buffered
saline,z and 1 ml of 10% sodium azide.
To this stock 5 gm of bromelin (Nutritional Biochemicals Corp.) was added for
incomplete Rh reagents. The diluted antisera was clarified by centrifugation and
frozen in 5 ml aliquots, which were
thawed for dispensing into Microtiter
plates. Once thawed, any unused residuum was discarded. To one drop of diluted antiserum one drop of red cell suspension was added and the plate was
incubated a t room temperature for eight
to ten hours. After being inclined at 45"
for a few minutes, the settling pattern
'PGL Paper 82. This work was supported by grant
GM 17173 from the U. S . National Institutes of Health.
The buffered saline was prepared from two stock solutions, one containing 9.46 gm NazHPOI/liter of 0.85%
saline, the other with 9.07 gm KHn/POl/liter saline. To
225 ml of the first solution and 75 ml of the second,
1700 nil of saline was added.
695
696
NEWTON E. MORTON AND MANABU YAMAMOTO
TABLE 1
ABO blood g r o u p s
~~
Population
0
A
B
AB
Total
303
134
68
39
4
68
74
9
32 1
164
154
81
10
73
138
17
45
23
91
52
1
33
18
5
33
12
46
32
0
16
24
2
702
333
359
204
15
190
254
33
699
958
268
165
2090
Test
Pingelapese
field
lab
field
lab
field
lab
lab
Mokilese
Ponapean
Kusaiean
Other
-
Total
All A
xz test
of panmixia
d.f. = 1
5.74
0.08
1.40
0.02
0.80
0.38
5.10
-
+ bloods tested in the laboratory were A1 t
TABLE 2
M N S blood groups
xa
Population
Test
Pingelapese
Mokilese
Ponapean
Kusaiean
Other
field
lab
field
lab
field
lab
lab
-
Total
Mss
Nss
MNss
MS-
NS-
21
9
27
19
0
14
24
6
466
209
160
86
6
71
122
13
194
108
111
60
7
76
66
12
1
0
12
1
12
3
14
22
120
1133
634
MNS-
8
4
Total
702
333
359
204
15
190
254
33
2090
0
1
1
1
0
15
25
1
35
16
1
13
16
1
16
93
94
ccEE
RZRZ
ccee
RoRo
Total
test
of panmixia
d.f.=2
0.04
2.07
1.45
3.47
2.16
1.68
9.72
TABLE 3
R h blood groups
Population
Test
Pingelapese
Mokilese
Ponapean
Kusaiean
Other
Total
field
lab
field
lab
field
lab
lab
-
CCee
R'R1
Ccee
RlRo
CcEe
RIR2
ccEe
R*RD
640
296
234
129
13
143
174
24
34
21
94
26
0
23
13
2
27
15
18
42
2
20
60
5
0
1
7
5
0
2
5
0
1
0
4
0
0
0
0
0
0
0
2
2
0
2
2
2
702
333
359
204
15
190
254
33
1653
213
189
20
5
10
2090
All bloods tested in the laboratory were D
x2 test
of panmixia
d.f. = 3
2.84
0.98
22.84
3.78
0.59
1.65
12.93
-
+ (Rhl + ).
was read with a hand lens or low-power
binocular microscope.
This Microtiter procedure uses only disposable plastic ware and is highly economical of reagents (Linbro MVC-96
plates are cheaper and are satisfactory
unless centrifuging is necessary). Anti-A,
anti-B, anti-M, and anti-N were diluted
1/160, a n t i 4 1/16. Anti-C, anti-E, and
anti-e werediluted 1/160 with bromelin
stock, anti-c 1/120. In the laboratory,
where the reagents can be kept frozen
at - 2 5 ° C for at least several months,
the Microtiter procedure is highly reliable. In the field repeated partial thawing due to inadequate refrigeration led
697
MICRONESIAN BLOOD GROUPS, HAPTOGLOBINS
TABLE 4
Haptog lobins
x 2 test
of panmixia
d.f. = 1
Population
Pingelapese
Mokilese
Ponapean
Kusaiean
Other
Total
1-1
2-1
2-2
Tot a1
HPO
233
23
29
44
10
155
104
78
143
16
26
82
82
60
7
414
209
189
24 7
33
0
1
0
5
0
0.00
1.38
2.05
6.54
339
4 96
257
1092
6
-
-
TABLE 5
x'
Test offield-lab homogeneity by factor and population
Factor
B
A
Population
M
N
S
C
C
e
E
Pingelap
0.53
0.08
1.99
0.14
0.68
0.29
1.36
0.37
0.47
Mokil
0.01
0.50
1.04
0.16
0.40
0.01
0.22
24.94
2.29
Microtiter system were detected in the
laboratory, but such errors must have
been rare in the field since heterogeneity
does not approach significance. Accordingly, we pooled field and laboratory tests
in every system and population, except
for the Rh field tests on Mokil, which
were rejected in deriving the gene frequencies of table 6 and in subsequent
analyses.
Even with exhaustive sampling there is
a tendency for x2 tests of panmixia to be
inflated because relatives are not independent samples, and this bias is exaggerated under field conditions where family
members are likely to donate blood together. Therefore we are not particularly
disturbed by the significant ~2 tests in
tables 1-4. The Kusaiean sample, which
in each rable shows a significant departure from panmixia, was nonsignificant
for isozymes (Yamamoto and Fu, '72).
to progressive loss of activity of anti-E
(Rh3), but other reagents remained
strong.
Haptoglobins were determined by starchgel electrophoresis of serum separated in
Honolulu and kept frozen at - 25 "C (Poulik, '64).
RESULTS
Excluding duplicate samples, 2090
bloods were typed (tables 1-3). There
were 1092 haptoglobin typings, omitting
six sera with such low activity as to be
unclassifiable (table 4).
To determine whether field tests were
reliable, we examined the 2 x 2 contingency tables of reaction ( or - ) X test
(laboratory or field) for each factor (table 5). Of the 18 tests, only anti-E on
Mokil was significant (P < < O.OOl), due
to loss of activity during field work. A few
false positive reactions with anti-c in the
+
TABLE 6
Gene frequencies assuming panmixia within populntions
ABO
Pingelapese
Mokilese
Ponapean
Kusaiean
Rh
MNS
Population
HP
A
B
0
Ms
Ns
MS
NS
RI
R'
R"+r
Hpl
0.300
0.335
0.280
0.393
0.056
0.222
0.130
0.085
0.644
0.443
0.589
0.521
0.177
0.257
0.296
0.247
0.809
0.650
0.625
0.667
0.005
0.045
0.014
0.013
0.009
0.048
0.065
0.074
0.951
0.799
0.871
0.829
0.022
0.115
0.059
0.128
0.027
0.086
0.071
0.043
0.750
0.359
0.360
0.468
Hpe
0.250
0.641
0.640
0.532
698
NEWTON E. MORTON AND MANABU YAMAMOTO
Low frequencies are found for the al- expected from the low frequency of shortleles B, S, RO, and R2, which on Pingelap range migration.
and Miokil appear to have been absent
LITERATURE CITED
among the survivors of typhoon Lengkieki
two centuries ago, and to have been in- Morton, N. E. 1964 Genetic studies of northeastern Brazil. Cold Spr. Harb. Symp. quant.
troduced by migrants (Morton et al., '72).
Biol., 29: 69-79.
Presumably the common Caucasian Rh
N . E., and R. E. Rosenfield 1967 A
negative allele has also been introduced, Morton,
new Rh allele, rvn(R - 1 . 2 . \ \ . 3 . W 4 ) . Transfusion,
but it was not distinguished from Ro
7: 117-119.
since all bloods tested in Honolulu were Morton, N.E., R. Law, I. E. Hussels and G . F.
Little 1972 Pingelap and Mokil atolls: historiRh positive (Rhl +). All group A and AB
cal genetics. Amer. J. Hum. Genet., 24: 277bloods tested in Honolulu were A1 .
289.
Pingelap has low gene frequencies of B Poulik, M. D. 1964 Starch gel-immunoelectrophoresis. Ann. N. Y. Acad. Sci., 121 ; 470483.
and S, and high frequencies of N, R', and
Hpr. This is not true for the neighboring Steinberg, A. G., and N. E. Morton 1973 Immunoglobins in the Eastern Carolines. Am. J .
atoll of Mokil. Despite their propinquity
Phys. Anthrop., 38: 699-702.
and known intermigration, leading to Wegmann, T. G., and 0. Smithies 1966 A simple hemagglutination system requiring small
spread of the gene for achromatopsia
amounts of red cells and antibodies. Transfrom Pingelap to Mokil, and of Gm',5s6
fusion, 6: 67-73.
in the reverse direction (Morton et al., Yamamoto, M., and L. Fu 1973 Red cell iso'72), the two gene pools appear to have
zymes in the Eastern Carolines. Am. J. Phys.
Anthrop., 38: 703-708.
differentiated almost independently, as
+
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