Anthropol.

Sci.

103(3),

Genetic

227-234,

1995

Polymorphism

New

ADA

of

Variants

FENG

1

JIN1*,SAITOU2,SUNS,HORAI5ofDepartment
411,

Taidong4

Department

of Evolutionary

Genetics,

411,

Taidong,

ADA

of

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Taiwan,

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theconversion

INTRODUCTION
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228

JIN et al.

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RESULTS
In
were

the

present

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besides

two

ADA
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alleles,

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ADA*

and
1

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The

IEF

ADA

Fig. 1Polymorphismin Taiwan

The sampling places of aboriginal populations

Aboriginal

Populations

229(Gaoshan) of Taiwan in the present study .

phenotype patterns of ADA are shown in Fig. 2 and those of schematic patterns areshown in Fig. 3. Because phenotypes ADA Taiwanl-1 and ADA Taiwanl-2 sharedbands with those of ADA 1 and ADA 2, respectively, they were considered to beheterozygotes of a variant allele ADA*Taiwan1 and common alleles ADA* 1 andADA*2, respectively (see Figs. 2 and 3). Phenotype ADA Taiwan 1 found in Amis
shared three bands with those of ADA Taiwan 1-1, and these shared bands did notmatch to any band of phenotype ADA 1. Therefore, phenotype ADA Taiwan 1 was

230

JIN et al.

Fig. 2 IEF patternsof ADA phenotypes

observed

ADA

1: lanes

1,5,8,11,12,14,16

and 23. ADA

2 and

18. ADA

Taiwan1:

lane

Fig. 3 Schematic

Taiwan1-1:

15. ADA

lanes

Taiwan2-1:

IEF patterns

in Gaoshan

populations .
lanes 4 , 6, 7, 20 and 21. ADA 2: lanes
3, 10, 13, 17 and 19 . ADA Taiwan1-2:
lane 9. ADA
lane 22.
2-1:

of ADA phenotypes

observed

in Gaoshan

populations.

ADA Polymorphismin TaiwanAboriginalPopulations

231

considered to be a homozygote of variant allele ADA*Taiwan1. Phenotype ADA
Taiwan2-1 was observed in a single individual. As in the case of ADA Taiwan11, it shared three bands with ADA 1, while the remaining band showed a distinct
IEF position different from any other phenotypes. Therefore, phenotype ADA
Taiwan2-1 was considered to be a heterozygote of a variant allele ADA*Taiwan2
and a common allele ADA*1.
In typing of IEF at pH 4-6.5, both variant phenotypes were quite similar to ADA
2-1 but seemed to be a little different. Thus, we retyped all the individuals of ADA
2, ADA 2-1, and suspicious ADA phenotypes by using IEF at pH 4.5-5.4 and 4.24.9. Seven phenotypes including ADA 1, ADA 2-1, ADA 2, ADA Taiwanl-1,
ADA Taiwanl-2, ADA Taiwanl, and ADA Taiwan2-1 were distinctively recog
nizedonly under the pH range 4.2-4.9 (see Fig. 2).
The phenotype and allele frequencies of the nine Gaoshan populations for the
ADA locus are listed in Tables 1 and 2, respectively. The observed values are ingood agreement with those of the expected (see Table 1). Allele frequency
distribution showed a considerable variation among the nine Gaoshan populations.
Frequencies of allele ADA*2 varied from low (0.032 for Yamis and 0.038 for
Tsous), intermediate (0.050 for Atayals, 0.057 for Bununs, 0.066 for Rukais, 0.074
for Puyumas, 0.087 for Saisiats) to high (0.125 for Paiwans and 0.139 for Amis).
Both Amis and the Paiwans possess the variant allele (denoted as ADA*Taiwan 1)with considerable frequencies (0.033 for Paiwans and 0.035 for Amis). We foundan individual of ADA*Taiwan 1 homozygote in Amis (see Table 1). One individualin the Saisiats was identified to have another new allele, ADA*Taiwan2.
DISCUSSION
There have been several reports on ADA variant alleles such as ADA*3, ADA*4,
ADA*5, and ADA*6 since its genetic polymorphism was recognized. All of theabove variants were recognized through the conventional starch-gel electrophoresis(Hopkinson et al., 1969; Dissing and Knudsen, 1969; Detter et al., 1970; Radamet al.,1974). To compare the new variants found in the present study with the known
variants, conventional starch-gel electrophoresis was also employed. The variant
phenotypes ADA Taiwan1-1, ADA Taiwan2-l, and ADA Taiwan 1 showed thesame electrophoretic mobility with that of ADA 1, while the variant ADA Taiwan 12 showed no electrophoretic mobility difference from the pattern of ADA 2-1. Therefore, two variant alleles ADA*Taiwan1 and ADA*Taiwan2 detected under the IEF
typing can be considered as subtypes of ADA* 1 under the starch-gel electrophoresis.
In contrast, none of the bands showed any similarity with published patterns underthe IEF typing (Harris and Hopkinson, 1976).
In the present study, we noticed that the two Gaoshan populations (Amis andPaiwans) had a relatively high frequency of ADA*Taiwan1 allele. There are so far

232

JIN et al.Table 1. ADA phenotype distribution in the nine Gaoshan populations of Taiwan*A

ADA

Table

2.

Allelefrequency

Polymorphismin Taiwan

Aboriginal(•}SE)

in theGaoshan

of

ADA

nine

Populations

populations

233

of

Taiwanno ADA variants possessing such a high frequency as ADA*Taiwan1 in the studies
of ADA polymorphism. It is of interest to note that the allele frequency of ADA*2
are also high (ca 0.13) in both Amis and Paiwans. In fact these two populations
showed the highest ADA*2 frequency among Asian populations reported thus far.
The frequencies of ADA*2 allele in the remaining seven Gaoshan populations
ranged from 0.032 to 0.087. These values are higher than those (0.010-0.023) inthe residents of Okinawa (Omoto, 1982; Omoto et al., 1975), but similar to those(0.030-0.081) in ethnic minorities of southern China (Li et al., 1989; Omoto et al.,1993). It is not clear whether the high frequency of both ADA*Taiwan1 and ADA*2
alleles in Amis and Paiwans indicates the genetic affinity of these two populations.
This could happen merely by chance.
Geographically, Taiwan is located on the sea shelf of the mainland China.
Paleontological evidences indicated that the Island was part of the Asian Continent
during the last glacial period, and the area was laid open for human settlement fromthe mainland (Bowles, 1977). The other geographically close area are the Luzon
Island of the Philippines and Ryukyu Islands of Japan. However, Ryukyu islanders
have been shown to be genetically not closely related with Gaoshan if we compare
several polymorphic loci (F. Jin et al., unpublished study). Therefore, Gaoshan,
aboriginal ethnic groups of Taiwan, probably migrated from either southern China
or Southeast Asia in the past. If it is the case, the variant allele ADA*Taiwan1 may
exist in some of populations of these area. Further studies are necessary to clarify
this point.
ACKNOWLEDGEMENTS
The authors thank the Gaoshan people of Taiwan who donated us their blood.
We thank two anonymous reviewers for their helpful comments. This study was
supported by Grants-in-Aid for Scientific Research (International Scientific
Research Program) and for Scientific Research on Priority Areas (Prehistoric
Mongoloid Dispersal) of Ministry of Education, Science and Culture (Japan) to S. H.

234

JIN et al.
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