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Blood genetic markers in Sri Lankan populationsЧreappraisal of the legend of Prince Vijaya.

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 76:217-225 (1988)
Blood Genetic Markers in Sri Lankan PopulationsReappraisal of the Legend of Prince Vijaya
N. SAHA
Department of Physiology, Faculty of Medicine, National University of
Singapore, Singapore 0511
KEY WORDS Sri Lanka, Sinhalese, Tamils, Muslims, Serum
proteins, Hemoglobin, Red cell enzymes, Electrophoresis, Isoelectric
focusing, Population genetics
ABSTRACT
Serum protein (haptoglobin types; transferrin and group-specific component subtypes); haemoglobin and red cell enzymes (acid phosphatase, esterase D, glyoxalase I, 6-phosphogluconate dehydrogenase, adenylate
kinase, and phosphoglucomutase (locus 1)(subtypes) were studied in the Sinhalese, Tamils, and Muslims of Sri Lanka. The allelic frequencies of all the
polymorphic systems were similar in these populations without any significant
differences. A close look at the present results and earlier investigations on 13
polymorphic loci controlled by 37 alleles did not reveal any genetic characteristics in the present-day Sinhalese population that are distinct from those in
the. Tamils of Sri Lanka. As such, genetic evidence linking the legendary
origin of the Sinhalese population to East India (Prince Vijaya) is lacking.
INTRODUCTION
The populations of present-day Sri Lanka
are extremely diverse, with the Sinhalese
constituting approximately 70% of the population. The remaining population includes
the Tamils, Muslims, and other minorities,
like the aboriginal Veddahs and the Burghers. Both myth and mystery surround the
origin of the Sinhalese, who claim to be descendants of the peoples of North-East India
more than 2,000 years ago, according to the
legend of Prince Vijaya (Geiger, 1964).There
have been immigrations from North-East India (present-day West Bengal and Bangladesh) to Sri Lanka for centuries. However,
Basham (1964)holds that the Sinhalese came
from the western coast of India.
The origin of the Sinhalese language has
been described as basically northwestern
with considerable northeastern influence
(Geiger, 1938) of the Indic languages and as
quite distinct from the Dravidian languages.
The Tamils originally came from Southern
India (presently Tamilnadu) and migrated to
Sri Lanka in the remote past, as evidenced
by archeological discoveries in the northern
part of the country, which was ruled by a
0 1988 ALAN R. LISS, INC
series of monarchies. There have also been
more recent Tamil immigrants from southern India, who came as workers during the
British rule of the subcontinent and settled
in Sri Lanka. Many of the Muslims of Sri
Lanka claim Arab ancestry; they came as
traders to the Malabar coast of India (present-day Kerala) (Fig. 1). A small group of
Burghers are descendants of Dutch and Portuguese who intermarried with the local
population.
The main population groups, Sinhalese and
Tamils, are quite distinct in their social habits, religion, and language-but intermarriage between the two groups has been going
on for generations. Questions about the origin of the Sinhalese have interested many
authors, because this Indo-European-speaking Buddhist population is thought to be as
culturally (and presumably biologically) distinct from the Dravidian-speaking Hindu
Tamils as from the minority remnants of the
Reeeived January 8, 1981; revision accepted September 24,
1987.
Address reprint requests to Dr. N. Saha, Department of Biochemistry, Faculty of Medicine, National University of Singapore, Singapore 0511.
218
N. SAHA
I
f
Bombov
\
R
Fig. 1. Different regions of India and Sri Lanka associated with the historical background of Sri Lankan
populations.
aboriginal Veddahs of Sri Lanka. However,
the Sinhalese system of kinship classification, cousin marriage, and the style of cooking, dress and housing are similar to those of
Dravidian-speaking areas of southern India.
Recent ethnographic work suggests that the
cultural differences between the Sinhalese
and Tamils and the Veddahs were overemphasized by earlier ethnographers (Brow,
1978).
Genetic studies on the populations in Sri
Lanka have been mostly limited to blood
groups and abnormal hemoglobins in the past
(De Silva, 1957; Wickremasinghe and Pon-
nuswamy, 1963; Wickremasinghe et al.,
1963). The first detailed study of genetic
markers in the S r i Lankan populations was
carried out by Kirk et al. [1962], followed a
decade later by a study on red cell enzymes
in the Sinhalese population (Roberts et al.,
1972). Kirk (1976) first tested the legendary
origin of the Sinhalese by genetic distance
analysis and concluded that the Sinhalese
are nearer to the Bengalis and the Tamils
than to the western Indians.
Since the study by Kirk et al. (1962), no
comparative study has been carried out
among the Sinhalese and Tamil populations
'Abbreviations: obs.
Groupspecific
components
Transferrin
1-1
2-1
2-1M
2-2
0
All
Haptoglobin
= observed; exp. =
22
IF2
IS2
2
2
All
IF
IFIS
IS
Y
c2-1
c2-2
C3-1
C3-2
c3-3
C1-B
C2-B
C1-D
C2-D
c1-l
22
Phenotypes
Systems
expected.
25.00
14.00
11.00
27.00
18.00
8.00
0
103.00
7.80
1.00
30.00
1.00
72.00
0
104.00
1.42
71.00
27.00
5.00
1.00
0
0
1.00
0
1.00
0
106.00
1.93
20.10
23.85
7.08
26.94
15.99
9.03
0.00
102.99
69.77
30.01
3.23
0.81
0.17
0.00
0.81
0.18
0.81
0.17
105.96
104.01
-
73.62
2.62
27.77
Sinhalese
No.
No.
obs.
exp.
Tamils
19.00
13.00
15.00
32.00
14.00
8.00
1.00
102.00
13.60
0
0
0
1.00
101.00
5.44
0
0
68.00
1.00
101.00
0.00
65.00
29.00
3.00
3.00
16.89
23.19
7.96
25.63
17.60
9.73
0.31
101.31
0.79
0.18
101.00
-
64.96
28.87
3.21
2.41
0.56
0.02
68.06
1.00
101.00
-
~
No.
- exp.
3.06
3.00
28.88
29.00
No.
obs.
13.00
13.00
7.00
5.00
7.00
2.00
0
47.00
2.75
10.30
15.92
6.15
7.50
5.79
1.36
0.00
47.02
0.76
0.21
46.97
-
-
49.00
49.00
1.36
29.00
13.00
3.00
0
1.00
0
0
0
1.00
0
47.00
5.03
27.58
15.32
2.13
0.76
0.21
36.00
1.00
12.00
35.00
14.00
0
Muslims
No.
No.
obs.
exp.
GC2
GC2
GC'F
GC'S
TfB
TfD
HP'
HP2
HPO
Allele
TABLE 1. Distribution of serum protein polymorphism in Sri Lankan populations'
-
0.4417
0.2621
0.2961
0.8113
0.1745
0.0047
0.0047
0.0047
-
0.1587
0.8413
-
0.4069
0.2794
0.3088
0.0049
0.0049
0.8020
0.1782
0.0149
0.1750
0.8250
0.0099
-
-
0.4681
0.3617
0.1702
0.0106
0.7660
0.2128
0.0106
-
0.1429
0.8571
Gene frequencies
Sinhalese
Tamils
Muslims
220
N. SAHA
of Sri Lanka. Several new genetic marker
polymorphisms have been described since the
work of the above authors and that of Roberts et al. (1972) on which no data on the Sri
Lankan populations are available.
Isoelectric focusing now makes it possible
to carry out a detailed study of some blood
genetic markers for population genetics, but
to date no study of isoelectric variations has
been reported in the Sri Lankan populations.
In view of the above developments and the
controversies over the origin of the major
Sinhalese population of Sri Lanka, we present here the results of a genetic marker study
in three Sri Lankan populations, Sinhalese,
Tamils, and Muslims.
MATERIALS AND METHODS
A total of 260 unrelated male Sri Lankans,
106 Sinhalese, 105 Tamil Hindus, and 49
Muslims, formed the study sample. They
were attending the Singapore Anti-Tuberculosis Association (SATA) Clinic for medical
examinations required for their employment
as foreign workers. Samples of blood were
collected in EDTA tubes. Plasma and packed
cells were separated immediately by centrifugation at 2,000 rpm for 30 minutes. Both
the plasma and the packed cells were stored
frozen at -80°C in aliquot portions. Hemoglobin and red cell enzymes, except the ones
mentioned below, were examined by horizontal starch-gel electrophoresis with standard
techniques. Serum proteins (haptoglobin,
transferrin albumin, and ceruloplasmin)
were studied by polyacrylamide gel electrophoresis by the Ashton and Braden buffer
system. Red cell acid phosphatase and esterase D were phenotyped with fluorogenic
stains. Red cell glyoxalase I polymorphism
was examined by the iodine staining method.
Serum transferrin was subtyped by isoelectric focusing at pH 3.5-10 and pH 5-7 after
desialation with neuraminidase. Group-specific component subtypes were identified by
isoelectric focusing at pH 4-6.5 using the
sulfosalisylic acid precipitation method. Red
cell phosphoglucomutase polymorphism was
studied by starch-gel electrophoresis and
isoelectric focusing at pH 5-7.
The gene frequencies of the different polymorphic loci were estimated by gene counting. The Hardy-Weinberg equilibrium was
calculated by the chi-square (x2)test.
RESULTS AND DISCUSSION
Serum proteins
The distribution of serum protein phenotypes and gene frequencies in the Sinhalese,
Tamils, and Muslims of Sri Lanka is presented in Table 1. No significant difference
in gene frequencies for Hp types, transferrin,
and group-specific component subtypes was
observed between these three populations.
The frequency of Hp' was 0.16 in the Sinhalese, 0.17 in the Tamils, and 0.14 in the
Muslims. Kirk et al. (1962) observed a similar frequency of Hp' (0.17) in the Sinhalese;
however, the Tamils of Sri Lanka in their
series had a lower frequency of Hpl (0.13)
compared to the frequency (0.17)observed in
the present study. An even lower frequency
of Hp' has been reported in the Tamils of
South India (Ananthakrishnan and Kirk,
1969; Ananthakrishnan, 1972). The frequency of Hp' in the Bengalis has been reported to be similar (Das et al., 1974;
Mukherjee et al., 1974; Papiha et al., 1975;
Saha, 1986). A solitary example of Hpo was
observed among the Tamils of the present
series. The incidence of Hpo has been reported to vary from 1%to 2% in Sri Lankan
populations (Kirk et al., 1962). The lower incidence of Hpo in the present series could be
due to the use of a sensitive polyacrylamide
gel electrophoresis technique in the present
investigation in place of the conventional
starch-gel electrophoresis used in earlier
investigations.
The frequencies of Tf and Gc subtypes were
also similar in these three groups. The frequency of Tfc' was found to be 0.81, 0.80,
and 0.76, respectively, in the Sinhalese, Tamils, and Muslims of Sri Lanka. A low frequency of Tf c 3 ,and T f Dwas also present in
these three groups. Only one case of Tp in
the heterozygous form was detected in the
Sinhalese. A similar low incidence of Tp
and T P in the Sinhalese and Tamils was
observed by earlier workers (Papiha and
Wastell, 1974). A much higher frequency of
Tf (0.87)was observed in the Bengali Muslims of Bangladesh (Saha, 1986), while a
lower frequency was reported in the populations of West Bengal (Reddy et al., 1984). A
similar frequency of T f ' has also been reported in the Tamils of Madras (0.77) and of
Singapore by Kamboh and Kirk (1983) and
Saha and Tan (1983).
The frequency of Gc (subtype) alleles also
did not exhibit any significant difference
among the Sinhalese and Tamils of Sri
Lanka. However, the Muslims had a significantly higher frequency of GcIS (0.36) than
that observed in the Sinhalese (0.26) and
Tamils (0.28). The Muslims also had a lower
frequency of Gc2 (0.17) compared to that in
the Sinhalese (0.30) and Tamils (0.31). The
89
50
30
10
90
2
18
36
56
58
3
1
62
1.37
12.25
27.37
40.99
56.04
2.92
0.04
59.00
28.55
7.90
0.55
37.00
0
15
26
41
57
1
1
59
28
9
0
37
1-1
2-1
2-2
A
AC
C
All
1-1
2-1
2-2
All
All
All
1-1
2-1
2-2
'Abbreviations: obs. = observed;exp. = expected.
Adenylate kinase
6-Phosphogluconate
dehydrogenase
Glyoxalase I
Esterase D
CB
All
B
4.28
33.44
65.28
0.00
103.00
52.96
41.07
7.96
101.99
8
26
69
0
103
53
41
8
102
A
AB
58
0
53
5
0
6
28
55
105
0
104
1
Acid phosphatase
104.01
1.97
0.01
106.00
104
2
0
106
A
AE
E
All
Haemoglobin
53.11
4.78
0.11
58.00
57.10
4.80
0.10
62.00
2.16
17.68
36.16
56.00
4.49
31.01
53.50
0.00
89.00
46.94
36.11
6.94
89.99
104.00
1.00
0.00
105.00
Tamils
No. obs.
No. exp.
Phenotypes
Systems
Sinhalese
No. obs.
No. exp.
26.00
0.00
0.00
26.00
26
0
26
0
15.02
0.97
0.02
16.01
1.00
8.00
16.00
25.00
2.57
16.61
26.81
0.75
46.74
36.67
15.66
1.62
54.00
53.00
1.00
0.00
54.00
15
1
0
16
0
10
15
25
3
16
27
1
47
37
15
2
54
53
1
0
54
Muslims
No. obs.
No. exp.
A@
AK~
PGD~
PGD~
0.1216
0.8484
0.9746
0.0254
0.1829
0.8171
0.2794
ESD~
GLO~
GL@
0.7206
E~D'
P"
Pb
0.2039
0.7961
0.00
0.0094
HbE
Pa
0.9906
0.0446
0.9569
0.9597
0.0403
0.1964
0.8036
0.2778
0.7222
0.2247
0.7753
0.00
0.0048
0.9952
0.0000
1.0000
0.9688
0.0312
0.2000
0.800
0.1759
0.8241
0.2340
0.7553
0.0106
0.0093
0.9907
Gene frequencies
Sinhalese
Tamils
Muslims
HbA
Allele
TABLE 2. Distribution of huemoplobin and red cell enzyme Dolvmorphisms in Sri Lankan uouulations'
222
N. SAHA
GcW frequency observed in the Sinhalese and
Tamils of Sri Lanka is much higher than
that reported in a limited number of Indian
populations (Walter et al., 1984).
i
I
I
I
I
dI
I
L
d
I
!
.!
!
r
I
1
I
4
I
d
I
7
I
,
,
I
5
2
n
G
P
:
Haxmoglobin and red cell enzymes
The distribution of haemoglobin and five
polymorphic red cell enzymes in the Sinhalese, Tamils, and Muslims of Sri Lanka is
presented in Table 2. A low frequency of HbE
has been observed in all three populations
without any significant difference. A similarly low frequency of HbE has been reported
in the Tamils of South India and in the Bengalis (Saha and Banerjee, 1973; Ghosh et al.,
1981). The Veddahs in Sri Lanka have been
reported to have a high frequency of HbE
(Kirk et al., 1962; Wickremasinghe et al.,
1963). Wickremasinghe and Ponnuswamy
(1963)found no HbE in more than 1,000 samples of Sinhalese, while De Silva et al. (1959)
had reported a high frequency in one locality
of Sri Lanka. The latter might have been due
to admixture with Veddahs in that area. An
important feature to note is that the incidence of HbE is very low and is present in
both the Sinhalese and the Tamils of Sri
Lanka. This suggests that HbE has originated with the Veddahs rather than being
introduced from South or East Asia, in which
case the incidence should have been greater
in the Sinhalese, contrary to Kirk's (1976)
suggestion.
The frequency of red cell acid phosphatase
alleles was also very similar among the three
populations of Sri Lanka. The frequency of
p a was found to be 0.20,0.22,and 0.23 in the
Sinhalese, Tamils, and Muslims, respectively. Roberts et al. (1972)reported a slightly
higher frequency of pa (0.25)in their series of
Sinhalese. No pc was detected in either the
Sinhalese or the Tamils. One example of AP
CB was encountered in the Muslims. A low
frequency of p" has been reported in the Sinhalese and Tamils by previous workers (Ananthakrishnan, 1972;Roberts et al., 1972).A
similar frequency of AP alleles has also been
reported in the Bengalis (Das et al., 1970,
1974; Saha, 1986).
The frequency of EsD' was also the same
in the Sinhalese and Tamils (0.721,while that
in the Muslims was 0.82. We have observed
a similar frequency of EsD' in Bengali Muslims (0.73) by (Saha 19861, while Papiha et
al. (1975) reported a EsD' of 0.77 in Bengali
Muslims.
The frequencies of GLO alleles were also
found to be similar in the three groups, and
the frequency of GLO' was 0.19 in the Sin-
223
GENETIC MARKERS IN SRI LANKANS
TABLE 4. Gene frequencies in Sinhalese, Tamils, and Bangalis (13 loci and 37 alleles)1
Sinhalese
Bengalis
Tamils
ABO
P
4
r
(0.1880)
(0.1650)
(0.6470)
(0.1760)
(0.1940)
(0.6300)
(0.1844; 0.2369)
(0.2070; 0.2634)
(0.608S; 0.4996)
(0.5780)
(0.4220)
(0.6170)
(0.3830)
(0.6537; 0.5603)
(0.3463; 0.4397)
(0.6500)
(0.08201
(0.0380)
(0.0130)
(0.2170)
(0.6339)
(0.0508)
(0.0122)
(0.0263)
(0.2768)
(0.6706; 0.6730)
(0.0792; 0.1315)
(0.0550; 0.0277)
(0.0341)
(0.1611; 0.1509)
0.1587 (0.1710)
0.8413 (0.8290)
0.1750 (0.114; 0.133)
0.8250 (0.886; 0.867)
0.1736 (0.175; 0.170; 0.185)
0.8264 (0.825; 0.830; 0.815)
0.8113
0.1745
0.0047
0.0047 (0.003)
0.0047 (0.003)
0.8020
0.1782
0.0149
0.0000
0.0049 (0.0006)
0.8710
0.1250
-
0.4417
0.2621
0.2964
0.0094
0.4069
0.2794
0.3088
0.0048 (0.004)
(0.752)
0.2521 (0.248)
0.0194 (0.01)
0.2039 (0.253)
0.7961 (0.734)
- (0.013)
0.7206 (0.763)
0.1829
0.0254 (0.016)
0.1216 (0.083); x2, 4.8
0.6968 (0.739)
0.2873 (0.261)
0.2247 (0.268)
0.7753 (0.730)
(0.001)
0.7222
0.1964
0.0403 (0.022)
0.0446 (0.063)
0.6810 (0.736)
0.3095 (0.264)
0.016
0.0048
MN
M
N
Rh
R1
RZ
Ro
r'
r
HP
HPl
H D ~
Pa
P"
PC
ESD'
GLO'
PGD~
AK2
PGM:
PGe
PGi@
PGi@
0.0040 (0.015; 0.012)
0.2403 (0.203; 0.199)
0.7519 (0.790; 0.801)
0.0078 (0.007)
0.7326 (0.771)
0.2077 (0.2127)
0.0250 (0.040)
0.0385 (0.087)
0.7286 (0.722)
0.2571 (0.272)
(0.003)
0.0048 (0.003)
'Gene frequencies in parentheses have been computed from the literature.
halese and Tamils and 0.20 in the Muslims.
A similar frequency of GLO' (0.21) had been
reported in the Bengalis and Tamils (Saha,
1986; Ghosh, 1977). The frequency of PGD'
was 0.04 in the Tamils and 0.03 in the Sinhalese and Muslims of the present series. A
similar frequency of PGD' had been reported
earlier in the Sinhalese, Tamils, and Bengalis (Roberts et al., 1972;Ananthakrishnan,
1972; Das et al., 1970, 1974; Papiha et al.,
1975; Saha, 1987).
The frequency of & was found to be 0.12
in the Sinhalese and 0.04 in the Tamils. Roberts et al. (1972)reported a frequency of 0.08
in their series of Sinhalese. A higher frequency of A3? (0.08) has also been reported
in the Tamils of South India (Ananthakrishnan, 1972). The higher A P frequency ob-
served in the Sinhalese in the present series
should be interpreted with caution since the
size of the sample was rather small.
Table 3 shows the distribution of PGM subtypes in the Sinhalese, Tamils, and Muslims
of Sri Lanka. The frequencies of the PGM
subtype were also found to be similar in these
three populations. The frequency of PGM1+
was slightly higher in the Sinhalese (0.55)
and Muslims (0.58)than in the Tamils (0.53),
while the frequency of other alleles was similar in the three groups. The frequency of
PG@+ was higher in the Sinhalese (0.02)
than in the Tamils (0.005) and Muslims. A
low frequency of PG@+ has been reported
in the Bengalis (Saha, 1986). One example of
a new variant @I 6.5) has been observed in
the Tamils and has also been reported in
224
N. SAHA
other Indian and East Asian populations
(Saha: unpublished observation).
Monomorphic system
Serum albumin, lactate and malate dehydrogenase, phosphohexose isomerase, phosphoglucomutase (locus 21, and superoxide
dismutase were monomorphic. The absence
of the lactate dehydrogenase variant is to be
noted, as this variation has been reported in
many populations of India, particularly in
Bengal. In the present investigation, however, it may be due to the small size of the
sample.
In Table 4 the gene frequencies of 13 loci
(37 alleles) in Sinhalese, Tamils, and Bengalis have been computed from our investigations and from the literature. It is clear
that the frequencies of these alleles in the
Sinhalese and Tamils of Sri Lanka are similar and that they are not significantly different from those in the Tamils of India or the
Bengalis. The population genetics studies
(present and earlier ones) so far carried out
in the Sinhalese have not revealed any significant differences in the gene frequencies
at 13 polymorphic loci from those of the Tamils in Sri Lanka. The allelic frequencies in
the Sri Lankan populations, in general, are
similar to those in other Indian populations.
Furthermore, the results of five Gm allotypes
and two Km allotypes show that the presentday Sinhalese are closer to the Sri Lankan
and Indian Tamils than to the Bengalis (Matsumoto et al., in press).
The present and earlier investigations produced no evidence whatsoever that the Sinhalese are genetically nearer to Eastern
Indian populations than to the Tamils or to
other South Indian populations. Even though
there has been some legendary connection of
the Sinhalese with East Indian or East Asian
populations through trade or social links,
there is no evidence to suggest that the present-day Sinhalese population is in any way
genetically distinct from the Tamils of Sri
Lanka.
This finding might be due to free intermarriage between the two populations for several generations in the past in spite of the
fact that they practice considerably different
religion, language, and social systems. As far
as we can see, the genetic evidence falls short
of supporting the legend that the Sinhalese
are descendants of Prince Vijaya.
ACKNOWLEDGMENTS
The author is grateful to Dr. N.C. Sengupta, Medical Director of the Singapore
Anti-Tuberculosis Association (SATA) for
permission to carry out this work. The technical assistance of the staff of SATA, Madame C.H. Ho and Jumiah bte Basair and the
secretarial help of Miss W.F. Chan are gratefully recorded. The project was generously
supported by grants from SATA, the Singapore Turf Club, the Shaw Foundation, and
the Singapore National Heart Association.
LITERATURE CITED
Ananthakrishnan, R (1972) Further studies on the distribution of some serum protein and enzyme groups in
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