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Distribution of HLA antigens in Kotas and Badagas of the Nilgiri Hills South India.

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 74:125-129 (1987)
Distribution of HLA Antigens in Kotas and Badagas of the Nilgiri
Hills, South India
A. SELVAKUMAR, B.K. MOHAN RAJ, C.DAMODARAN, AND
P. CHANDRA SEKHARAN
Forensic Sciences Department, Madras-600 004, India
KEY WORDS
Tribes, Population study
ABSTRACT
Blood samples from 103 Kotas and 58 Badagas residing in the
Nilgiri Hills, South India, were examined for HLA-A and -B antigen profiles.
The Kota group was characterized by fairly high frequencies of A2 and B7
antigens as well as the haplotype A2-B7. The frequencies of Aw19, A28, and
Bw22 were found to be higher in Badagas than in Kotas. The results are
compared with the literature available on other Indian populations.
The Nilgiri Hills area, popularly called
“Blue Mountain,” covers an area of 984
square miles in the Nilgiri District (Tamil
Nadu State) situated in the southwest of the
Indian peninsula. The percentage of tribal
population in this region is the highest in
Tamil Nadu and includes the endogamous
groups Kotas, Badagas, Todas, K m m b a s ,
Irulas, and Paniyas (Ghosh, 1973; Verghese,
1974). The artisan-musician Kota tribe were
one of the earliest inhabitants of the Nilgiris,
living on the slopes between 4,000 and 6,000
feet. The entire Kota population now distributed in seven villages had lived in isolation
from other tribal groups for centuries. The
Kotas, closely resembling the Todas, however, appear to have originated as a plains
tribe of the neighbouring Kerala coast, as
shown by anthroposcopic, anthropometric,
and genetic distance studies (Ghosh et al.,
1977). Badagas, on the other hand, are relatively recent migrants from the adjacent
Karnataka. They are agriculturists and form
the single largest community in the Nilgiri
Hills (John Kumar, 1984).
The HLA system, the most polymorphic
and complex set of genetic markers known
in man, is of valuable significance in anthropological studies (Dausset, 1981). Distribution of HLA antigens in various Indian
population groups has been reported (Ftaha,
1975; Bale et al., 1980; Mittal et al., 1982;
Verma et al., 1983; Pitchappan et al., 1984).
However, data on Indian tribal populations
is rather scanty (Papiha et al., 1983). Therefore it was felt worthwhile to investigate the
0 1987 ALAN R. LISS. INC.
distribution of HLA antigens in Kotas and
Badagas of the Nilgiri Hills, South India.
MATERIALS AND METHODS
An HLA typing laboratory was set up at
the Government Arts College, Udhagamandalam, the district headquarters of the Nilgiris. One hundred three unrelated Kotas
were selected from five Kota villages-Kollimalai, New Kothagiri, Kilkothagiri, Trichigadi, and Kunda Kothagiri; 58 unrelated
Badagas were from Udhagamandalam, Aravenu, Horasholai, and Dimbatty. Figure 1
shows the locations of the sample collection
in the Nilgiri Hills.
Five milliliters of venous blood (in heparin,
50 IU/ml) was drawn in a sterile tube from
each individual; the samples were then
transported cold (24°C) to the laboratory
within 3 hours. For HLA typing, the heparinised blood was diluted with an equal volume of phosphate-buffered saline (PBS),
layered on Lymphoprep (Nyegaard, Norway),
and centrifuged at 1,800 rpm for 20 minutes.
The interphase lymphocytes were collected,
suspended in PBS, and washed twice (centrifugation, 1,000 rpm, 10 minutes) to remove
the contaminating nonlymphocytic white
cells. The pure lymphocyte suspension was
adjusted to 2,000,000 cells/ml. HLA-A and
-B typing was performed by NIH two-stage
microlymphocytotoxicity assay (Ray et al.,
1976) with an NIH (USA) starter Kit (containing 83 sera) and five sera of local origin.
Received November 17, 1986; revision accepted May 6,1987.
126
A. SELVAKUMAR ET AL.
Fig. 1.Location of Kota and Badaga villages (Nilgiris,
India).
These local sera (r value .6 to .9) were earlier
collected from multiparous women (Madras)
and first screened by us for HLA antibodies
(Selvakumar et al., 1987). Their specificities
were also compared with NM sera and confirmed by two well-established tissue typing
laboratories at New Delhi (AILMS) and Madurai (Madurai Kamaraj University).
The NIH starter kit and the local sera covered 12 specificities in HLA-A locus (Al, A2,
A3, A9, Aw24, A10, A26, A l l , Aw19, Aw31,
Aw32, Aw33) and 17 specificities of HLA-B
locus (B5, B7, B8, B12, B13, B14, B15, Bw16,
B17, B18, Bw21, Bw22, B27, Bw35, B40,
Bw41, Bw51). The phenotype frequency PF),
gene frequency (GF), standard error of gene
frequency (SE GF), two-locus haplotype frequency @F), coefficient of linkage disequilibrium (A), and chi-square with Yate's correction were calculated by 2 x 2 table as described by Mittal(1976).
RESULTS AND DISCUSSION
The distribution of HLA-A and -B antigen
phenotypes and their frequencies in Kotas (n
= 103) and Badagas (n = 58) are shown in
Table 1.In Kotas significantly increased frequencies of A2 (P < lov6; PX24 < lop5)and
B7 (P <
P x 24 <
were observed.
Badagas possess higher frequencies of A3 (P
< .025), Awl9 (P < .001), A28 (P < .005),
B8 (P < ,051, B12 (P < .005), B13 (P < .Ol),
B15 (P c .OW, Bw22 (P <
and Bw51
( P < .05). However, when the P value was
multiplied by 24 (the number of specificities
listed), the higher frequencies of Awl9 (Px24
< .024), A28 (Px24 < .048), and Bw22
(Px24 < lo-*) only were still significant.
These findings show that while A2 and B7
occur more frequently in Kotas, Aw19, A28,
and Bw22 do so in Badagas. The incidence of
A26 is found to be very low in Badagas and
totally absent in Kotas.
Table 2 gives the HLA gene frequencies in
Kotas and Badagas along with the reported
data in other Indian tribal and nontribal populations. It is evident that the frequency of
A2 in Kotas is the highest (45.63%)followed,
but not closely, by the tribal Koyas (23.40%)
of Andhra Pradesh, South India (Papiha et
al., 1983); in other Indian groups the A2 frequency is only in the range of 10.30-21.10%.
The frequency of B7 allele in other Indian
populations is between 3.45 and 18.48% only
whereas in Kotas it is extremely high
(45.63%). It is hence evident that the Kotas
are characterized by the high frequencies of
HLA-A2 and B7. The allelic frequency of
Awl9 is the highest in Badagas (12.07%)
closely approached by the population of the
plains of South India (11.58%); in others the
range is 2914.44%. HLA-A28 gene is, however, found to be fairly uniformly distributed
in most of the Indian groups including Badagas (5.17-7.10%) except in the secluded tribals, Kotas (0.49%) and Koyas (1.10%). As for
the Bw22 allele, it is distinctly higher in
Badagas (9.48%)than in others (0.49-2.50%).
Incidentally, it is also of interest to note that
A26, either absent or present in very low
127
HLA IN NILGIRIS TRIBES, INDIA
TABLE 1. HLA-A and -B antigens i n Kotas and Badayas (i'rilgiris, India)'
HLA
antigen
A1
A2
A3
A9
A10
All
Awl9
Aw24
A26
A2B2
AB5
337
B8
B12
B13
B14
B15
B17
B18
Bw22
B27
Bw35
B40
Bw5 1
B-
Kotas (n = 103)
%GF
n - %PF
7.77
16
15.53
45.63
94
91.26
6.80
14
13.59
5.34
10.68
11
2.91
5.83
6
15.05
30.10
31
2.91
5.83
6
1.94
3.88
4
0
1
0.49
0.97
11.16
23
7.28
14.56
15
94
45.63
.~ 91.26
1.46
2.91
3
3.40
6.80
7
2.43
4.85
5
2.91
5.83
6
3.88
7.77
8
2.43
4.85
5
1
0.49
0.97
0.49
0.97
1
1.94
3.88
4
6.80
7
3.40
11.17
22.33
23
0.49
0.97
1
26
SEGF
n
1.94
4.71
1.82
1.61
1.91
2.70
1.19
0.97
11
20
16
10
6
16
14
4
1
6
12
8
4
5
13
9
4
10
4
2
11
5
7
14
3
0.49
1.88
4.71
0.84
1.28
1.09
1.19
1.37
1.09
0.49
0.49
0.97
1.28
2.33
0.49
Badagas (n = 58)
%PF
%GF
SEGF
18.97
34.48
27.59
17.24
10.34
27.59
24.14
6.90
1.72
10.34
13.79
6.90
8.62
22.41
15.52
6.90
17.24
6.90
3.45
18.97
8.62
12.07
24.14
5.17
17
9.48
17.24
13.79
8.62
5.17
13.79
12.07
3.45
0.86
5.17
10.36
6.90
3.45
4.31
11.21
7.76
3.45
8.62
3.45
1.72
9.48
4.31
6.03
12.07
2.59
14.65
X<
2.86
3.86
3.45
2.73
2.11
3.45
3.23
1.72
0.86
2.11
0.61
55.16
5.76
2.05
1.85
0.02
13.18
1.49
2.70
10.26
2.44
1.72
1.93
3.11
2.59
1.72
2.73
1.72
1.22
2.86
1.93
2.28
3.23
1.49
0.01
107.37
3.91
9.82
6.74
0.37
4.38
0.81
2.97
20.12
2.60
2.05
0.21
4.72
Pvalue
NS
<
Px24*
< 10-5
< ,025
NS
NS
NS
NS
< ,001
NS
NS
< ,005
< ,024
< ,048
NS
110-6
< .05
< ,005
< .01
NS
< .05
NS
NS
10-5
NS
NS
NS
NS
< 10-4
NS
NS
NS
< .05
NS
'PF, phenotype frequency;GF, gene frequency.
'Assigned after studying the cell reaction in three duospecific sera.
* P value, multiplied by the number (24) of specificities listed.
frequency (0.86%)in the two populations now
studied (Kotas and Badagas, respectively),
occurs more frequently in other Indian populations (2.90-4.30%).
The haplotype frequencies and the linkage
disequilibria figures are given in Table 3. In
Kotas significant positive linkage disequilibrium was seen for two haplotypes, A2-B7
(t= 10.74) and A3-B7(t=3.22), and significant
negative linkage disequilibrium for three
haplotypes, A2-B5(t = -2.09), A2-B40 (t=
-2.21), and All-B7 (t= -2.97). On the contrary, the Badagas possess no haplotype with
significant positivelnegative linkage disequilibrium, though the haplotype A2-Bl5 occurs more frequently (35.43960; t =1.06).
Further, the Badagas show contrast with the
Kotas by the total absence of the two haplotypes A2-B7 and A3-B7, the former of
which is dominantly characteristic of the Kotas. Mittal et al. (19821, Verma et al. (19831,
and Pitchappan et al. (1984) have reported
haplotype frequencies in Indian populations
after HLA typing including A2, A3, All, B5,
B7, and B40. Indian Hindus (Mittal et al.,
1982)and North Indians (Verma et al., 1983)
do not possess any of the haplotypes A2-B7,
A3-B7, A2-B5, A2-B40, or All-B7. However,
the haplotypes A337 and A2-B40, each with
significant positive linkage disequilibrium,
have been observed in the population of the
South Indian plains (Pitchappan et al., 1984).
It is hence interesting here to note that while
the Kotas show a significant negative linkage disequilibrium for A2-B40, they constitute, as reported now, the second Indian
population to show significant positive linkage disequilibrium for A3-B7.
The Kotas have been studied extensively
for many non-HLA genetic markers (Ghosh,
1973; Ghosh et al., 1977); there is, however,
no report in the literature on the genetic
constitution of the Badagas except for the
study on sickle cell trait (Lehmann and Cutbush, 1952). Ghosh et al. (1977) list as the
distinctive characteristics of the Kotas the
extremely low frequency of the A1 gene and
the total absence of A2 in the ABO system, r
(cde) in the Rh system, LDHCa'-' among the
enzyme system as well as of abnormal hemoglobins. Now, based on the present study on
HLA, it is evident that the Kotas are further
128
A. SELVAKUMAR ET AL.
TABLE 2. HLA gene frequencies (%) in Kotas, Badagas and other Indian populations'
HLA
allele
a
Kotas
(n = 103)
A1
A2
A3
A9
A10
All
Awl9
Aw23
Aw24
Aw25
A26
A28
A29
Aw30
Aw31
Aw32
Aw33
Aw34
Aw36
AB5
B7
B8
B12
B13
B 14
B15
Bw16
B17
B18
Bw21
Bw22
B27
Bw35
B37
Bw38
Bw39
B40
Bw41
Bw42
Bw44
Bw45
Bw47
Bw48
Bw49
Bw50
Bw51
Bw52
Bw53
Bw54
Bw55
Bw57
Bw58
Bw60
Bw61
Bw62
Bw63
B-
7.77
45.63
6.80
5.34
2.91
15.05
2.91
NT
1.94
NT
0.00
0.49
NT
NT
0.00
0.00
0.00
NT
NT
11.16
7.28
45.63
1.46
3.40
2.43
2.91
3.88
0.00
2.43
0.49
0.00
0.49
1.94
3.40
NT
NT
NT
11.17
0.00
NT
NT
NT
NT
NT
NT
NT
0.49
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
12.60
h
b
Badaeas
(n = 58)
Kovas'
(n 94)
d
Hindus3
(n = 138)
e
South India
(n = 385)
f
North India
(n = 233)
g
West India
(n = 118)
East India
(n = 92)
9.48
17.24
13.79
8.62
5.17
13.79
12.07
NT
3.45
NT
0.86
5.17
NT
NT
0.00
0.00
0.00
NT
NT
10.36
6.90
3.45
4.31
11.21
7.76
3.45
8.62
0.00
3.45
1.72
0.00
9.48
4.31
6.03
NT
NT
NT
12.07
0.00
NT
NT '
NT
NT
NT
NT
NT
2.59
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
14.65
3.70
23.40
5.30
NT
NT
28.70
NT
5.90
13.30
0.50
4.30
1.10
0.00
0.00
0.00
0.50
NT
NT
NT
13.30
NT
8.00
4.30
NT
3.20
1.60
1.10
1.10
25.50
2.20
0.50
0.50
1.60
2.70
0.00
NT
NT
5.30
0.50
0.50
5.30
4.80
NT
NT
NT
.~
NT
1.60
3.20
1.10
NT
NT
NT
NT
NT
NT
NT
1.10
24.30
20.60
10.30
7.10
14.00
2.90
14.50
NT
0.40
13.60
0.00
2.90
6.00
1.50
1.80
1.10
2.90
12.00
0.40
0.00
4.90
14.90
3.70
3.70
10.70
3.30
0.40
8.30
2.20
14.00
2.60
0.70
1.10
0.70
17.00
3.30
1.80
0.40
9.10
1.10
0.00
10.30
0.70
0.40
0.70
0.00
0.70
7.50
6.40
0.70
0.40
0.70
12.40
1.50
2.60
1.80
3.30
3.30
1.40
13.10
16.41
6.30
16.72
3.44
16.57
11.58
NT
NT
NT
NT
6.58
NT
NT
NT
NT
NT
NT
NT
8.89
18.14
10.12
1.84
5.34
3.03
0.13
5.89
0.39
10.12
0.52
2.77
2.50
0.91
10.27
NT
NT
NT
8.12
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
17.57
11.60
21.10
7.40
10.40
1.10
9.50
3.30
1.50
3.70
1.30
3.30
7.10
3.50
2.40
1.70
4.80
0.90
NT
0.60
4.80
16.60
9.70
5.50
8.00
3.10
0.60
8.50
0.40
9.00
2.60
2.20
2.40
3.90
4.40
1.70
0.20
0.20
14.30
0.60
0.20
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
0.20
NT
NT
NT
NT
NT
NT
5.70
15.12
13.15
6.12
14.13
3.45
13.15
8.44
0.43
13.15
0.00
3.01
7.03
NT
NT
NT
NT
NT
NT
NT
19.41
6.57
13.64
2.14
6.57
2.14
0.85
4.77
1.28
7.49
5.67
1.28
1.71
0.85
13.64
NT
NT
NT
8.87
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
22.53
36.41
16.30
4.35
11.96
16.85
14.13
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NR
23.91
18.48
20.11
22.83
14.67
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
NT
~NR
~~
C
la and b, present study; c, Papiha et al. (1983); d, Mittal et al. (1982);e, Pitchappan et al. (1984);f, Verma et al. (1983);g, Bale et al.
(1980); h,Raha (1975).NT = not tested. NR = not reported.
ZTribalgroup of Andhra F'radesh, South India.
3North and South Indian Hindus in USA.
129
HLA IN NILGIRIS TRIBES, INDIA
TABLE 3. H e
a, and t values o f HLA-A and -B haplotypes
in Kotas and Badagas ( l o )
Poaulation
Hadotvae
Kotas
A1-B7
A2-B5
A2-B7
A2-Bl5
A2-B40
A3-B7
A9-B40
All-B7
All-B40
Aw19-B7
A1-Bw22
A2-BI2
A2-Bl5
A2-B40
All-B15
Badagas
HF~
il
46.31
0
704.43
-11
0
0
70.94
21.67
0
19.70
12.81
28.87
18.37
35.43
15.75
27.56
-76.16
208.25
-42.52
-89.81
20.87
14.66
- 148.64
0.58
-8.16
20.17
-5.86
17.93
- 10.17
13.62
t value'
0.50
-2.09
10.74
-1.55
-2.21
3.22
1.55
-2.97
0.05
-0.52
1.33
0.33
1.06
0.55
0.87
'Haplotype frequency.
't values greater than 2 indicate significant positive linkage disequilibrium and those less than -2
indicate significant negative linkage disequilibrium.
characterized by the extremely high frequency of A2 and B7 antigens and also of the
haplotype A2-B7. As yet, however, no plausible explanation could be offered for the
uniqueness observed in the Kotas except the
suggestion of selective advantage.
ACKNOWLEDGMENTS
The authors acknowledge the excellent assistance from the tribedgeneral population
and from personnel belonging to voluntary
bodies, universities, and government departments. Special thanks go to Prof. J.I.V. Jayapaul and his students and t o Dr. M.
Sethuraman (Department of Medical Anthropology, Tamil University, Udhagamandalam). The gift of HLA sera by NIH (USA)
is duly acknowledged.
The financial assistance to A. Selvakumar
and B.K. Mohan Raj by the Bureau of Police
Research and Development, New Delhi, is
thankfully acknowledged.
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badagas, kotas, distributions, hla, antigen, indian, south, hill, nilgiri
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