Result and Discussion
4.1 Genotype Frequencies
4.2 Gene Frequencies
4.3 Chi-Square test for departure from Hardy-Weinberg expectations
4.4 Pair-wise comparison of gene frequencies between population
4.5 Expected heterozygosity & Average heteozygosity
4.6 Gene diversity between populations
4. 7 Genomic affinities among populations
4.8 Comparison with other Indian Populations
4.9 Comparison with world populations
4.10 Gene Flow Among Populations
Chapter IV
I
85
FiJ;u r 4.1 f>hotograph howing garo e gel electrophoresi ( 1.5%) lu 0 I in ertion/dclction fn~gm nt si~:c
Lane 6 homozygous for the - (-I-) alleles Lane 8 homozygous for the - (+/+) alleles Lane 9 homozygous for the - (+/-) alleles Lane 7 Marker Ladder Product Size : +/+ = 650bp -1- = 350bp
Lane 9 (+I·) --
Lane 8 (+/+) --Lane 7 Marker Ladder
Lane 6 (-/·)
Figure 4.2 Photograph showing Agarose gel electrophoresis (1.5%) Alu PLAT insertion/ deletion fragments sizes
Lane 5 homozygous for the - (+I-) alleles Lane 6 homozygous for the - (-I-) alleles Lane 7 Marker ladder Lane 10 homozygous for the - (-I-) alleles Product Size : +/+ = 400bp -1- = 11 Obp
Lane 10 (·/-) ----
Lane 7 Marker Ladder Lane 6 (+/+) ---~ Lane 5 (+/-)-------<
86
Fi~ure 4. Photograph showing Agaro. e gel electrophoresis (LS-%) E R ~ locu primer ntnplificd Toq I digested D A ' u ,es
Molecular size marker Pvu/1 digested 0 X 17 4 RF DNA Lane 4 homozygous for the = +/+Alleles Lane Marker Ladder Lane 9 heterozygous for the = +/- alleles Lame 17 homozygous for the = -1- alleles Product Size +/+ band = 1350 bp -1· band = 1350 bp, 900bp, 450bp. +I· band = 900 bp, 450 bp.
Lane 9 (+/·) ----
Lane 7 Marker Ladder
Lane 17 (-/·) ---Lane 4 (+/+) ----
Figure 4.4 Photograph showing Agarose gel electrophoresis (1.5-%) LPL 2.112.2locus primer amplified Taql digested DNA size
Molecular size marker Pvu/1 digested 0 X 17 4 RF DNA. Lane 13 heterozygous for the = +I- alleles Lane 2 heterozygous for the = +/+ alleles Lane 3 heterozygous for the = -1- alleles Lane 7 Marker ladder Product Size : +/+band= 171bp, 148 bp. +/-band= 319bp, 171bp, 148bp. -1· band = 319bp
Lane 13 (+/-)----
Lane 7 Marker Ladder
Lane 3 (-/-) ---Lane 2 (+/+) ----
H7
Ia hie 4.1 (;enol)lll.' fn"quendes al dghl Alu inserlion/ddelion pol~·morphil.' lud in three Jrih<'' of Bu,tar, ( 'hhattisgarh
I l'upulalions
i I - -
Ahujhmarill Bison-horn Maria Muria
( ~)(.:LI'\ ( icnotypc (icnolypc (icnolypc
I I I I 1- -I- 11 I I I tl- -1- 11 I/+ 1)- -1- 11 - --- --
Al111111 Nl I(' X I 'I 2) 52 X 25 21 58 X 22 I 'I 49 \\ u 15.38 36.54 48.08 100 1180 43.10 43.10 IIXJ _1623_ 44.90 38.77 100
- -~ -"-- --~- --Al11 Ai'O 21 21 6 52 32 23 4 59 2'1 12 8 49 Ill) 44.23 44.23 1.54 100 54.24 3X.48 6.78 100 59.18 24.49 16.33 100
-- ---~------ ~ -- ---- - - ---- ~- ----- -- --AI11ACE 15 29 10 54 28 24 8 (i) 9 34 6 49 uu 27.78 53.70 1852 100 46.67 40.0 13.33 100 18.37 69.39 12.24 IOU
Alii Cll4 49 5 0 54 57 3 0 (i) 48 I 0 49 % 90.74 9.26 () 100 95.0 5.0 0 100 97.96 2.04 0 100
----
Alulll 13 17 23 53 7 17 35 59 10 14 2) 49 (% 24.53 32.07 43.40 100 11.87 28.81 59.32 100 20.41 28.57 51.02 100
Alui'XIIIB 2) 21 4 50 36 21 2 59 32 13 4 49 % 50.0 42.0 8 100 61.02 35.59 3.39 100 65.31 26.52 8.16 100
Alu PV92 16 23 12 51 14 32 12 58 15 21 13 49 % 31.30 45.10 23.53 100 24.14 55.17 20.69 100 30.61 42.86 26.53 100
Alu PLAT 22 24 7 53 25 23 12 60 17 26 5 48 % 41.51 45.28 13.21 100 41.67 38.34 20 100 35.41 54.17 10.44 100
n =number ofindividual
Table 4.2 Genotype frequencies at seven unlinked Restriction fragment length polymorphic (RFLP) loci in three tribes ofBastar, Chhattisgarh
Populations
Abujhmaria Bison-horn Maria Moria
L<lcus Genotype Genotype Genotype
+I+ +!- -1- n +I+ +/- -I- n +I+ +!- -I- n
ESR 19 27 7 53 12 35 14 61 25 21 13 49 % 35.85 50.94 13.21 100 19.67 57.38 22.95 100 51.02 42.85 26.53 100
NAT 24 26 4 54 22 33 5 60 36 11 2 49 % 44.44 48.15 7.41 100 36.67 55.0 8.33 100 73.47 22.45 4.08 100
PSCR 8 18 20 46 I 25 34 60 3 22 24 49 % 17.39 39.13 43.48 100 1.67 41.67 56.66 100 6.12 44.90 48.98 100
T2 22 22 10 54 13 32 13 58 13 21 15 49 % 40.74 40.74 18.52 100 22.41 55.18 22.41 100 26.53 42.86 30.61 100
LPL 15 31 6 52 20 32 8 60 14 26 9 49 % 28.85 59.61 11.54 100 33.33 53.34 13.33 100 28.57 53.06 18.37 100
ALB 14 29 II 54 19 26 15 60 II 28 10 49 % 25.93 53.70 20.37 100 31.67 43.33 25.0 100 22.45 57.14 20.41 100 CYPIA 16 26 12 54 21 28 10 59 16 18 13 47 % 29.63 48.15 22.22 100 35.59 47.46 16.94 100 34.04 38.30 27.66 100
.. n lllilllberofmdiVIdual ------
HH
.·\ total or I 1>.1 unrelated i ndi' iduah li·orn t hrcc tri hcs, vi1. A hu j hmaria (A B M ), Bi sonlwrn
\I aria ( BIIR) and Muri;~ (MliR) w.:r<: g<:notyp.:d li>r 15 hialldic polynwrphic loci: J:ight <trt:
1 ""'rt ion/d.:lc:t ion - I ndc Is (mtN ll (·.A Ju ACF. Alu A 1'0. A lu FX II lB. A lu I> I. A lu (' D4, Alu
1'1 AI. Alu I'V'I2) and sen:n an: unlinked RFI.I's (I·:SR-I'vu/1. Nt\T-J(pnl. l'SCR-lillfl.
( · Y I' I A· Mspl. T2- Msp I. 1.1' l.-l'vu/1 and t\ I.B-1 lae I I 1). ( 'hromosomal locations of these
marKers arc given in chapter! V- Subjects and Methods. These markers arc core sd of
I lN t\ markers selected by the Department of Biotechnology ( DBT). Government of India as
a part of its I Iuman Genome Diversity (JIG D) initiative.
4.1 (;cnotypc Frequencies
The observed genotype frequencies in three tribes ofBastar, Chhattisgarh for eight
insertion/deletion polymorphic loci arc presented in Table 4.1. Of the total 72 Alu marker
groups, the frequency ofheterozygotes is more than 50% (the maximum frequency expected
liH any hi allelic locus) in only four marker-groups (for Alu ACE 53.70% in Abujhmaria and
69.39% in Muria, for Alu PV92 55.17% in Bison-horn Maria and for Alu PLAT 54.17% in
Muria). One locus in ABM and BHR and two loci in MUR showed excess heterozygotes
than expected for a biallelic locus. Homozygotes for the 'absent' (-)allele of Alu CD4 are
absent in all tribes under study.
The observed genotype frequencies for seven autosomal unlinked RFLPs loci among
three tribes of Bas tar under study are given in Table 4.2.0f the total 63 RFLP marker
groups, in 9 marker-groups, the frequency ofheterozygotes is more than 50%- the maximum
frequency expected for any biallelic locus. Three loci inABM (ESR, LPL & ALB), four loci
in BHR (ESR, NAT, T2 and LPL) and two loci in MUR (LPL and ALB) showed excess
heterozygotes than expected for a bialleilic locus. The frequecy ofheterozygotes in LPL
locus is more than 50% in all the tribes under study. Two RFLP loci in Muria, three RFLP
loci in Abujhmaria and four RFLP loci in Bison-horn Maria showed excess heterozygotes
than expected for a biallelic locus.
Table 4.3 Allele frequencies at eight Alu insertion/deletion polymorphic loci in
Abujhmaria, Bison-horn Maria and Muria tribes ofBastar, Chhattisgarh
Abujhmaria Bison-horn Maria Muria
Locus p(+) :!;SE q(-) N p(+) ;tSE q(-) N p(+) +SE q(-) N
A1u mt NUC (+) 0.3365 0.046 0.6635 104 0.3534 0.044 0.6466 116 0.3878 0.049 0 6122 98
A1u APO (+) 0.6635 0.046 0.3365 104 0.7373 0.041 0.2627 116 0.7143 0.046 0.2857 9X
ALU ACE(+) 0.5463 0.048 0.4537 108 0.6667 0.043 0.3333 120 0.5306 0.0450 0 4694 98
A1uCD4H 0.9537 0.018 0.0463 108 0.975 O.Ot4 0 025 t20 0.9898 0.010 0.0\02 98
A1u 01 (+) 0.4057 0.048 0.5943 106 0 2627 0.041 0.7373 118 0.3469 0.048 0.6531 98
A1u FXlllll (+) 0.7100 0.045 0.2900 100 0.7966 O.o38 0.2034 118 0.7857 0.041 0 2143 98
Alu PV92 (+) 0.5392 0.049 0.4608 102 0 5172 0.046 0.4828 116 0.5204 0.050 O.t7%i9X
Alu PI.Ar (+) 0.6415 +-- _ ____;
0.046 0.3585 106 0 niJOO 0 045 0.40110 t211 () 62)0 11.114'!11. (l\'1-=i i 9(1
' N numha of chromo:.-omcs. +- mscrt1on.-- deletion: SF:' Standard error
I
H9
Tuhlt• ~..1 Allele frequendt·s at 'ewn unlinked RFI.I' pul~·nwrphil: loci in Ahujhmaria, llhnn-hnrn Mnriu and Muriatriht•s of Hnstar, < 'hhuttisgarh
Ahujhmaria IJi\tJrt-lwrn Maria Muriu
t) .!_SE q(-) N Wl +SE q( ) N p(+) +SE q( ) N - -- _:_ ~---
I \]{ 0 t d 12 () 0-17 0 lXtlX I!Jh O..IHltt 0 ll-15 () ~16-1 122 (} 520·1 {) (j<:j() () 1796 'IX -
\ \I 0 f ,X 52 0 0-1' 0 li·IX lOX () (l.f 17 () ().j.l o l5K l 120 O.lH69 () 0 ~(J 0.15.11 <JX ---
{{)h {) {)){) 0.6.\0-\ 92 0.225 o.O.lX 0. 775 120 li.2X57 O.ll4h 0.7141 'IX ------· ---- ---- ___ :c;_ f--C. --- ---- -- ---- -- ------ ------- -----1'\l I\ ll. l
Ill {) 0-17 0.3XX9 lOX 0.5 004h_ 1--- 0.5 - 116 () 4796 0.050 _115201_ 'JX --- --- -·-- ---- - --- ·-· I 2 0.6
X65 0 U-IX {).41 l5 10-1 06 () 045 0.4 120 0.551 0.05{) {) 44'1 <JX - - - --- ------1----
11'1 0 5
\I I! 0.5 27X IUI4X 04722 I OX {) 5.1.l.l 0.046 0.4667 I 20 0.51 02 0.051 0.4X<JX '!X - --- ·--- ---
(\']'It\ 05 J7 U.04X 0.463 lOX 0.5932 0.045 0 406X IIX 0.5319 () ()53 0.4hX I 94
N number or chromosomes~+= insertion~-= deletion; SE =Standard error
~.2 Gene Frequencies
Table 4.3. shows the allele frequencies at eightAlu insertion/deletion polymorphic loci
with their standard errors in three tribes ofBastar under study. The Alu deletion allele'-' at
the Alu CD4 locus is human specific. At Alu CD4locus, Chimpanzees, Gorillas, Orangutans
and Gibbons are monomorphic for the'+' allele. The frequency of Alu CD4+allele varied
from about 1.02% in MUR to 4.6% in ABM.The frequency of this allele is about 2.5% in
BHR. The insertion allele,'+' at mtNUC is also human specific and reported to have occurred
before the divergence of human populations. The frequency ofmtNUC allele varied from
about 33.6% inABM and 38.7% in MUR. The frequency of this allele is about 35.34% in
BHR. Though the values of'+' allele are higher at some loci, the range of variation is low.
(< 0.1) atAluAPO,Alu CD4, Alu FXIIIB,Alu PV92 &Alu PLAT. Alu mt NUC,AluACE
& Alu Dlloci showed greater range of variation.
Table 4.4 shows the allele frequencies at seven autosomal unlinked RFLP polymorphic
loci in the populations under study. Among seven RFLP loci, the frequency of'+' allele
(insertion or restriction site-present') is less than 40% at one locus; PSCR, in three tribes
under study. The amount of variation is low ( < 0.1) at the three loci (LPL, ALB and CYP I A).
At one RFLP locus (PSCR) the upper limit of the range of'+' allele is 50% or less in all the
populations under study. The ESR locus ofBHR & T2 locus of MUR also show the upper
limit of the range of'+' allele 50% or less. At six other loci, the lower limit of the range of
'+'allele is more than 50% (NAT, LPL, ALB & CYPlA in all tribes under study; T2 in
ABM & BHR and ESR in ABM & MUR).
All the loci, insertion/deletion and RFLPs, are polymorphic in ABM. BHR and MlJR.
Figures 4.5, 4. 7 and 4.9 show the allele frequencies at IS biallelic polymorphic loci in ABM.
BHR and MUR respectively.
Figure 4.5 Allele frequencies at 15 biallclic polymorphic loci in Abujhmaria tribe of Bas tar, hhattisgarh
ALB 04722
0.5278
LPL 0 4135
[] 0 .5885
T2 0.3889
0.8111
PSCR 0.6304
0.3896
CYP1A 0.463
0.537
NAT 0.3148
0.6852
~ ESR 0.3888
Q) 0.6132 ~ ... nl
PLAT 0.3585
:E 0.6415
<t z CD4 c 0.9537
0.4537 ACE 0.5463
APO 0.3365
0.6635
01 0.5943
0.4057
FXIIIB 0.29
0.71
PV92 0.4608
0.5392
Alu mtNUC 0.6635
0.3365
0 0.2 0.4 0.6 0.8 1
Allele Frequency
90
1.2
~ Cl) ~ ... tU
:iE <( z c
Figure 4. 7 Allele frequencies at 15 biallclic polymorphic loci in Bi on-horn Maria tribe of Bas tar, Chhatti5garh
ALB 0.4667
[] 0 5333
LPL 0.4
0.6 p
12 0.5 0.5
PSCR 0.775
0.225
CYP1 A 0.4068
0.5932
NAT 0.3583
0.6417
ESR 0.5152
0.4848
PLAT 0.4
0.6
CD4 0.9537
ACE 0.3333
0.6667
APO 0.2627
0.7373
01 0.7373 0.2627
FXIIIB 0.2034
0.7966
PV92 0.4828
0.5172
Alu mt NUC 0.6466 0.3534
0 0.2 0.4 0.6 0.8
Allele Frequency
91
1.2
tn .... Q) .X .... ns
::E <( z c
Figure 4.9 Allele frequencies at 15 biallelic polymorphic loci in Muria tribe of Bastar, hhattisgarh
LPL
12
PSCR
CYP1A
NAT
ESR
PLAT
CD4
ACE
APO
01
FXIIIB
PV92
9 0.551
~ ~
Alu mt NUC
0 0.5 1 1.5
Allele Frequency
92
'
93
'llthle -4.5 The ( 'hi-squan· ,·aiUl'S (I d.f.) fur departure fmm liard~·-Wcinherg cxpt'Ctatiuns in 1 'i hialldil- pulnnurphil-lnl·i fur the data presented in 'litblcs -4.1 & -4.2
S.'lu. l.uci AHM MlJR IUIR - -·
I Alum! Nlll' 1.72 0.145 0.19 ------- ----
l Alu PV'I2 0.436 0.98 0.64 ---- -- - - ----- ----
\. Alul·XIIlll 0.02 2.2! 0.26 - ..... --·· --~·~--- .. --~- ------
4. AluD! 5.94 6.689 3.87 .-- - ---· ---- -- -------- --·- --· ----------
). AluAI'O ().()05 7.84 0.002 - - --- -------. ....
(>. AluAC'E 0.375 7.57 0.6 ---- ----·~·
7. Alu CD4 0.079 0.0052 0.04 -- -- --
8. Alu PLAT 0.295 1.!6! 2.29 -----
'1. ESR 0.289 0.98 1.35
!0. NA:r 0.727 0.88 2.31 ·-
11. CYPIA 0.054 4.37 0.02
12. PSCR 1.18 0.49 2.28
13. T2 1.102 0.98 0.62
14. LPL 2.73 0.26 0.74
15. ALB 0.323 1.01 1.01
4.3 Chi-Square test for departure from Hardy-Weinberg expectations
Table 4.5 shows the values of r.' tests for goodness of fit with Hardy-Weinberg
expectation for 15 biallelic polymorphic loci among three tribes under study. Six of 45 r.' tests for goodness of fit were significant (p<O.OS). All the populations showed significant
departure at locus Alu Dl. Alu APO, AluACE and CYPlA are significant in MUR only.
Departure from Hardy-Weinberg expectations is more than expected by random chance in
Muria tribe is more than ABM and BHR. About 25% of the loci are expected to show
significant departure by random chance, when level of significance is set at 0.05 probability.
But in ABM and BHR 6.67% loci (1/1 5) showed significant departure. In MUR 26.6% loci
showed significant diparture.
Table 4.6 The Chi-square values (ldf) for pairwise comparison of gene
frequencies between the populations
S.No. Loci ABMvsBHR ABM vs MUR BHRvsMUR
1. AlumtNUC 0.138537 1.150699 0.539867 2. AluPV92 0.210791 0.14183 0.004358 3. AluFXIIIB 4.412023 3.005199 0.077129 I
4. AluDl 10.20583 1.57289 :1.417904 j
-----
5. AluAPO 2.882458 1.213482 0.2R:'i)O:i -- -
7.
X
<).
Ill.
II.
12.
Ll.
14.
15. .
:\lu:\( 'I·
:\ 1u C ll4
A1u Pl.AI
LSR Ni\1
CYP I;\
PSCR - ---- - -------12
. ~- .. --1.1'1.
. --------ALB
(>.'1~5095
1.52:\~)5
!UC24 1'1
7.540564
0.%1 H25
1.450:l54 ..
10.62554 - ------
5.586826 -----------
0.084\9 f----~-
0.0\3808
h>r df \ Chi- square (0.05) = 3.84
0.1 0 llJ2lJ IUM>-177
4.735657 IJ1215X
0.1 JXI6:1 0.2H0519
3.575572 0.551 OO:l
14.X2tn:l 23.26492 -. . . - .. --- -
0.010139 1.543052 -- ·- - -- ---------- -- . . -- ---
3.03887H 2.107309 ---
--~ --~---- --7. I 75247 0.176918 -----
0.5\8648 \.06200\
0. I 275\3 0.230769
4.4 Pair-wise comparison of gene frequencies between population
The x' values for the pair-wise comparisons of gene frequencies at 15 biallelic
polymorphic loci between the populations (ABM vs BHR, ABM vs MUR,and BHR vs MUR)
arc given in Table 4.6. Of the total 45 pair-wise comparison of gene frequencies between
!he populations under study, II (about 24.4%) are statisfically significant.Most of the
significant comparisons are between Abujhmaria and Bison-horn Maria. Six of the fifteen
(about 40%) significant differences are between these groups at the loci AluFXIIIB, Alu D I,
Alu ACE, ESR, PSCR & T2.Abhujmaria shows three significant differences (20%) with
Muria (at loci Alu CD4, NAT & T2).8ison-horn Maria & Muria differ at two loci Alu ACE
& NAT. Thus Muria shows greater similarity with Bison-horn Maria than with
Abujhmaria.Abujhmaria shows greater difference with Bison-horn Maria.
Among 15 loci, AluACE, NAT & T2 loci showed significant differences in two of the
three possible comparisons. These three loci, together can differentiate all the populatios
under study.
Table 4. 7 Heterozygosity at individual loci based on eight Alu insertion/deletion
polymorphic loci in each of three tribes of Bas tar, Chhattisgarh
Locus Abujhmaria Bison-hom Maria Muria
AlumtNUC 0.4465 0.4570 0.4748
AluAPO 0.4465 0.3874 0.4081
AluACE 0.4957 0.4444 0.4981
AluCD4 0.0713 0.0487 0.0202
AluDl 0.4822 0.3874 0.4531
AluFXIIIB 0.4118 0.3339 0.3367
AluPV92 0.4969 0.4994 0.4992
AluPLAT 0.4485 0.4800 0.4687
95
Figure 4.6 IJ cterozygos ity at 15 biallclic polymorphic loci in Abujhmaria tribe of Bustur, hbattisgarh
0 0.1 0.2 0.3
Heterozygosity 0.4
0.4984
0.4972
0.5 0.6
96
Figure 4.8 Heterozygosity at 15 biallclic polymorphic loci in Bison-horn Maria tribe
of Bastar, hhattisgarh.
0.49778222
0.5
0.48262752
0.49953792
~ .... .---------------------------~~------------------------~ ~ ... ns
:::e <( IL.a..lo.:L::l
z c
0 0.1 0.2 0.3
Heterozygosity 0.4
0.44442222
0.49940832
0.5 0.6
97
Figure 4.10 Heterozygosity at 15 biallclic polymorphic loci in Muria tribe of Bas tar, hhattisgarh
0.4998
0.4948
0.4992
0.498
0.4992
0.4991
0 0.1 0.2 0.3 0.4 0.5 0.6 Heterozygosity
9H
Tahl .. -I.H llt'll'nl/) gusil)' al indi\·idualluci han'tl un sewn unlinkedl'\'stricliun fragment
knglh pul) ruurphic ( IU'LI'jluci in thl'\'e tribes uf Has tar, ( 'hhallisgarh
l.oru' Abujhmaria Hison-hurn Maria Muria
ISR O..J7.J.J 0.4995 0.4991
NAI 0.4114 0.4598 0.2593
!'SCI{ 0.465'1 0.3487 0.4082 ------- --- ----
-~ --- --
1'2 0.4753 0.5 0.4992 --- - ------·------- -- -----
LI'L 0.4850 0.48 0.4948 ----- ----·-------
/\Ill 0.4984 0.4978 0.4998 - -- ------
CYI'IA 0.4973 0.4826 0.4977 ----------
Table 4.9 Average heterozygosity and Standard error (SE) based on 15 biallelic
pulymorphie loci in each ofthree tribes ofBastar, Chhattisgarh -- --Tribe Average heterozygosity + SE
Abujhmaria 0.448009 0.026510
Bison-horn Maria 0.423362 0.030651 r-
Muria 0.425526 0.034320
4.5 Expected heterozygosity & Average heteozygosity
Table4.7, 4.8 and 4.9 & Figures 4.6, 4.8 &4.10 show the calculated heterozygosity at each of
the locus and over all loci among three tribes ofBastar. The heterozygosity values are higher at all loci
(>30%) with the exception ofAlu CD4 in all the tribes & NAT for MUR. The heterocygosity at the Alu
CD4 varied between 2.02% in MUR to 8.83% inABM. For NAT locus, the heterozygosity varied
between 25.93% in MUR & 45.98% in BHR. Maximum heterozygosity (>49%) was observed at the
lociAluACE and CYP1A(ABM & MUR),AluPV92 &ALB (ABM, BHR & MUR), ESR& T2
(BHR & MUR) and LPL (MUR). T2locus ofBHR shows maximum heterozygosity (50%).The
average heterozygosity over all loci varied from about 42% in BHR & MUR and about 45% in ABM.
Table 4.10 Results of gene diversity analysis for individual loci and for all loci considered
jointly
Locus 8, Hs CST
AlumtNUC 0.460369 0.459458 0.001979
AluAPO 0.415923 0.414021 0.004571
AluACE 0.486813 0.479421 0.015185
AluCD4 0.052857 0.052418 0.008307
AluDI 0.447792 0.440904 0.015382
AluFXIl!B 0.360502 0.357536 0.008229
AluPV92 0.498689 0.498501 0.000378
AluPLAT 0.470151 0.469569 0.001 ~38 ·-
~~
I Sl{ O..l'itJ'I4X o. 49 I 000 0.011967
'\AI OJ'I'I II 0 O.JXJ522 0.039057
J'S(' R 0.41-lhhO 0.40763 I 0.0 I 6952
!"~ 0.49X IT!. 0.491494 0.01.1405
I .I' I. 0.4X7465 0.4X6611 0.001752
Al.ll 0.49XX70 0.49!1676 (). 0003 89 -------· ----
C'YJ>IA 0.494161 0.492618 0.003122 .. --
All loci 0.432166 0.428225 0.009117 -------·
111
Total genomic diversity among the populations
II Diversity between rndividuals within populations '
G..,1
- Between population genomic diversity
4.6 Gene diversity between populations
Table 4.1 0 presents the results of gene diversity analysis for each locus and also for
all loci together among three populations under study. The total genomic diversity (H,) varied
0.360 (Alu FX!IlB) to 0.498 (ALB) with the exception of locus Alu CD4, for which the
value is 0.053. However, most of the genomic diversity is attributable to diversity between
individuals within populations (Hs). The percentage of genomic diversity attributable to
total genomic diversity (Gsrl varies between 0.03% (Alu PV92) and 3.90% (NAT).
Considering all loci taken together, only 0.91% ofthe total genomic diversity is attributable
to between populations.
Table 4.11 Genetic distance in three tribes of Bas tar, Chhattisgarh
Tribe ABM (1) BHR (2)
BHR (2) 0.0041 -
MUR (3) 0.0035 0.003 5
4. 7 Genomic affinities among populations
Table 4.11 shows the pair-wise genetic distances between the populations under study
calculated using the allele frequencies. Figure 4.11 shows the neighbour joining tree
constructed from this distance matrix. The three tribes group into two clusters. BHR &
MUR are the closest related. ABM is distinct from BHR & MUR.
r--------------BI son hon Mttr i;,
~-------------------------------------------------Abujl~ria
Figure 4.11 Neighbour joining tree depicting genomic relationships among three
tribes of Bastar, Chhattisgarh
100
Tahlt• .1.12 llricf description of twcnt}' six populations of India (locations of
, 11 rnplin~. sample si:r.c (n), lin~uistic affiliation and occupation) taken for compnrison
indudin~ 11rcscnt study ,., --· -· -----
'lrilllll 1•upulntiun I ,Ol'lltiun uf Samplin" "' Lingui~tk Affilialion Occuruttion - ---- -· ---- --- -- - ---- . -· ------- .
\hllilllll.lllil Jl;i'>[.tt 1)1\J'>IUil, ( hb.ll(l',!.!,tllh ;.1 I >r:tvJdiaH ShiJlmg cult•vatJtlll, hunting - ·- ·- 1---
BJ-.tlll-hlllll ~l.uJ:t IJa,lat I h\ l'>illn. < 'hhatli..,gcu h 6U l>ravtdtmt Ag,rl~.:ulturi:-.l, lluntLT- gathering ---- --- - --- r--- ----- ------- --------
fV1tHI.l l~a ... tar I)J\ J\Jon. < 'hhattJ'>garh 4'1 l>ravitlJall Agm:ulturist. l I untmg-gathcrml:!-- - ---- ---- - - -
llalh.t McJd!J)a 11Jadc:-.h 4X lmlo-l~unlpcan Agricultun! - - ·-· -------
1\.amar Madh)a 11radc ... h 57 l>raviJian h1od gatherer. Ba..,kdary - -- --- ---'- -
( '!m1da Bhnnpa Matlltya l'radesh 25 \ndo-l·:uropcan Agnculturc ---
( 'haukhulla IJhun_1ia Madhya flwJcsh 39 lndo·l~uropcan 1\gm:uhurc
Munda Sundcrgarh, Orissa 52 Austro-/\siatic Primarily agricultural laborers
Mcdinipur, West Bengal
l.odlw Mcdintpur, West Bengal 64 Austro-Asiatic J>rimarily agricultural laborers
Santa! Mcdinipur District 48 Austro-Asiatic Primarily agricultural laborers
East Bengal
Ka thana Tharu Kheri, UttarPradesh 60 Indo-Aryan Agriculturist, hunting and fishing
Ran a Tharu Kheri, UUarPradesh 60 Indo-Aryan Agriculturist, rearing pigs, fO\vls
Danguria(B) Babratach, UttarPradesh 60 Indo-Aryan Agriculturist, hunting and fishing,
Danguria Balrampur, UttarPradesh 60 Indo-Aryan Agriculturist, hunting and lishing
Pachchimaha Mahar~jgang, Uttar Pradesh 60 Indo-Aryan Agriculturist, rearing pigs, goats
Toto Jalpaiguri, West Bengal 30 Austro-Asiatic Primarily agriculturist
Mizo M1zoram 30 Austro-Asiatic Primarily agriculturist
Ho East Indian Region 54 Austro-Asiatic Primarily agriculturist
Tipperah Various location around 45 Tibeto-Burman Agriculturist
Agartala. Tripua.
lladaga Hilly tracts and valleys of 51 Dravidian Hunter gathers, now practice
Nilgiri Hills Tami!Nadu shifting or settled culti vatlOn
lrula Hilly tracts and valleys of 50 Dravidian Hunter gathers, now practice
Nilgiri Hills Tami!Nadu shifting or settled cultivation
Kota Hilly tracts and valleys 45 Dravidian Hunter gathers, now pract1ce
ofNilgiri Hills Tami!Nadu shifting or settled cultivation
Kurumba Hilly tracts and valleys 54 Dravidian Hunter gathers, now practice
ofNilgiri Hills TamilNadu shifting or settled cult1vatmn
Toda Hilly tracts and valleys 50 Dravidian Hunter gathc.rs. now practice
ofNilgiri HillsTamiiNadu shifting or settkd culti'.-ation
Koya Dora East Godavari District. 63 Dravidian Agriculturi'\t
Andhra Pradesh
Konda Reddy East Godavari District. 62 Dravrdian Agriculturi'\t
Andhra Pradesh - -- .
101
T~thlt• 4.1.' Allde frequencies 111 eight Alu insertion/deletion pol~'m11rphic l11ci in twcnt
,j, trilniiJlnpul~ttinns in lndi11 includin~t prt-scnt study -- -.----- ,-- -----c-·---- -----
Triht· Alu mt Alu Alu Alu Alu Alu Alu Alu
Nll(' AI'O ACE ('1)4 Dl FXJIIH l'V 92 I'LAT
J•(+) I'(+) p(+) p (-) p(+) II(+) p (+) p(+) i
-- ---
I i\bujlunmia 0.336 0.663 0.546 0.953 0.405 0. 710 0.539 0.641
i '-- ---- --------- -----------~
Bison-horn Maria OJSJ O.TJ7 0.666 0.975 0.262 0.796 0.517 0.600 - --
Muria 0.3X7 0.714 0.530 0.989 0.346 o.ns 0.520 0.625
llalha 0.625 0.691 0.646 0.094 0.427 0.698 0.563 0.574 -- ------ -- ---~ -
Kamar 0.463 0.649 0.640 0.018 0.342 0.746 0.554 0.535 --------- f-----
( 'hinda l~hlmjia 0.420 0.481 0. 796 0.038 0.620 0.731 0.407 0.556 -~---
Clmukatia Bhu11jia 0.513 0.622 0.649 0.090 0.289 0.750 0.412 0.729
Munda 0.587 0.300 0.640 0.080 0.320 0.667 0.481 0.660
Lodha 0.452 0.453 0.859 0.016 0.281 0.823 0.532 0.625
San tal 0.478 0.761 0.521 0.025 0.292 0.725 0.563 0.417
Katharia Tharu 0.416 0.847 0.585 0.031 0.411 0.791 0.696 0.814
RanaTharu 0.509 0.735 0.622 0.009 0.490 0.773 0.801 0.792
Danguria(B) 0.500 0.754 0.661 0.018 0.332 0.703 0.711 0.644
Danger 0486 0.900 0.75 0.042 0.337 0.815 0.770 0.710
Pachchimala 0.406 0.872 0.711 0.031 0.363 0.614 0.625 0.702
Toto 0.683 0.850 0.567 1.000 0.333 0.944 0.933 0.033
Mizo 0.426 0.845 0.603 1.000 0.286 0.648 0.862 0.586
Ho 0.453 0.819 0.731 1.000 0.452 0.802 0.565 0.713
Tipperah 0.459 0.863 0.590 0.012 0.313 0.846 0.811 0.549
Badaga 0.625 0.784 0.460 0.020 0.230 0.461 0.436 0.551
Iru!a 0.574 0.570 0.750 0.040 0.600 0.640 0.449 0.550
Kota 0.567 0.767 0.622 0.000 0.589 0.878 0.300 0.659
Kurumba 0.538 0.583 0.806 0.000 0.528 0.694 0.713 0.704
Toda 0.204 1.000 0.469 0.050 -i
0.300 0.806 0.255 0.406 -- - -·
KondaReddi 0.410 0.611 0.717 1.000 0.320 0.500 0.603 0.596
KoyaDora 0.427 0.672 0.777 0.974 0.409 0.800 0.654 '
0.644 i --
102
Tuhle -'·1-' The numht-r of individuals at eaeh loeus studied in twenty six
l"'l'uh<tions in lndi11 eumined for eomparison including present studv . ... ------· - ---
Trih<• Alum! Alu Alu Alu Alu Alu Alu Alu
NIJC AI'O ACE Cl>4 ()J FXIIIB J>V 92 PLAT I
1 Name n II n II n n II n --- ... ---- ' ----- -- ·-· -- ' ----··-- -
I
I I ., . J\ >llillll<Ula 52 52 54 54 53 50 51 53 .. ---. ----·· ---- ---···-··
llison-horn Maria 5K SK 60 60 59 59 58 60 --- .. - - -------- ---- ----
Mtu·ia 49 49 49 49 49 49 49 48 ·-·· -- --- ---- ---- --
llalha 4K 47 48 48 48 48 48 47 - ---
Kamar 54 57 57 57 47 57 56 57 ----- .
Chinda Bhlu1_jia 25 27 27 26 25 26 27 27 ----
Chaukatia Bhu11_jia 39 37 37 39 38 38 34 35
Munda 46 so 50 52 50 48 52 50
Lodha 64 64 64 64 62 62 64 64
San tal 46 46 48 40 48 40 48 48
Katharia Tharu 36 36 35 35 34 36 33 36
RanaTharu 53 53 53 53 53 53 53 53
Danguria(B) 48 59 59 55 59 59 59 59
Danguria 38 38 38 38 35 35 37 38
Pachchimaha 48 47 45 47 44 48 48 47
Toto 30 30 30 30 30 29 30 30
Mizo 27 29 29 26 28 27 29 29
Ho 53 53 54 54 52 53 54 54
Tipperah 74 80 78 82 80 78 74 82
Badaga 48 51 50 50 50 51 47 49
lrula 47 50 50 50 50 50 49 50
Kota 45 45 45 45 45 45 45 44
Kurumba 53 54 54 54 54 54 54 54
Toda 49 50 49 50 50 49 49 48
KondaReddi 56 54 53 54 50 50 53 57
KoyaDora 55 58 40 59 55 55 55 59 ..
n- number of mdJVJdual
103
Tahlc 4.1 S llclcnu~ ~osily at ind i\·idualloci hased on ei~ht Alu insertion/deletion
luci in'" cnty six populations of India including present study -
Trihc Alu rnl Alu Alu Alu Alu Alu Alu Alu
NIIC AJ'() ACE ('()4 I> I FXIIIB PV92 I'LAT i . ---- - -- ---- -
t 1\btqhmaria 0.4465 0.4465 0.4957 0.0883 0.4822 0.4118 0.4969 0.45<)<) - - --- - ----- ------- -- --
Bl..,on-horn Maria 0.4570 0.387.1 0.4444 0.0487 0.3873 0.3240 0.4994 0.4800 --- - --- -- ~- -- ----
Muria 0.4748 0.4081 0.4981 0.020 I 0.4531 0.3367 0.4991 0.4687 - - - --- -- -- ------- ------
llalha 0.4687 0.4270 0.4573 0.1703 0.4893 0.4215 0.4920 0.4890 - ----
Kamar 0.4972 0.4555 0.4608 0.0353 0.4872 0.3789 0.4941 0.4975 ~- ----·
Chinda llhunjia 0.4872 0.4992 0.3247 0.0731 0.4712 0.3932 0.4827 0.4937 -----
Chaukatia Hhunj ia 0.4996 0.4702 0.4555 0.1638 0.4109 0.3750 0.4845 0.3951 -
Munda 0.4848 0.4200 0.4608 0.0000 0.4352 0.4442 0.4992 0.4488
Lodha 0.4953 0.4955 0.2422 0.0314 0.4040 0.2913 0.4979 0.4687
Santa! 0.4990 0.3637 0.4991 0.0487 0.4134 0.3987 0.4920 0.4862
Katharia Tharu 0.4811 0.2591 0.4904 0.0276 0.4862 0.3294 0.4247 0.2964
RanaTharu 0.5000 0.3978 0.4697 0.0372 0.4935 0.2939 0.2939 0.3294
Danguria (B) 0.5000 0.3707 0.4481 0.0357 0.4366 0.4172 0.4101 0.4584
Danguria 0.4996 0.1800 0.3750 0.0821 0.4473 0.3005 0.3538 0.4113
Pachchimaha 0.4824 0.2227 0.4107 0.0617 0.4627 0.4737 0.4687 0.4183
Toto 0.4330 0.2550 0.4838 0.0237 0.4300 0.2605 0.3065 0.4951
Miw 0.4890 0.2619 0.4910 0.0000 0.4442 0.1572 0.4910 0.0638
Ho 0.4955 0.2563 0.4787 0.0000 0.4084 0.4561 0.2379 0.4852
Tipperah 0.4966 0.2364 0.3932 0.0000 0.4953 0.3175 0.4915 0.4092
Badaga 0.4687 0.3386 0.4968 0.0392 0.3542 0.4969 0.4918 0.4947
Irula 0.4890 0.4902 0.3750 0.0768 0.4800 0.4608 0.4947 0.4950
Kota 0.4910 0.3574 0.4702 0.0000 0.4841 0.2142 0.4200 0.4494
Kurumba 0.4971 0.4862 0.3127 0.0000 0.4984 0.4247 0.4092 0.416 7
Toda 0.3247 0.0000 0.4980 0.0950 0.4200 0.3127 0.3799 0.4823
I !
- -- -KondaReddi 0.4840 0.4753 0.4058 0.0000 0.4352 0.5000 0.4 78--l ! 0 -lX 11
Koya Dora 0.4894 0.4405 0.3459 0.0495 .)_
0.4834 0.3200 0.4522 0-l:iX.J ----- - -
104
Tahle 4. J(, ("hi square \'a lues (I df) for the pair-wise comparison of Gene
fn·qul·ndes of Ahhujmaria with twenty three populations of India -- - - - - -- - - - ----
i Popuhttion Alu rnt Alu Alu Alu Alu Alu Alu Alu
Nll(' AI'<) ACE {"I)" I)( FXIIIB PV92 PLAT ---- -- ------ ------- r------ ~-- ---
ll:tiiM ll .121117 () l41262 4.1 X7115X ]() .1.4 J.12 0 197XI II. 116 77 09 0.226436 1.91179114
-- -----+--------K.unar 7.05%21 0.101)29 4.0:W)69 3X'J.5721 0 097X36 0.699211 0 094376 5 137077
-- -----~ --
( 'hJJlda Bhunj1a 2.034821 9.87471 X 1924871 360.8794 12 95085 0.148763 4.937385 2.2114059
~----
( 'h.tu~atia llhunjia II 46189 11.6514.17 3.H21X29 279.65114 5.465028 0 696667 5.283983 2.948873
-----
Munda 46 5066 35.99065 2.457223 27X.4115 2.189139 0.56690 I 0.934723 0.101879 ----
l.odha 4.484376 14.44527 35 15591 295.665 5.468245 5.258521 0.016396 0.093781
-
Santnl 5.421441 2.852221 () 171181 241.0117 3. 772883 0.063 0 149217 13.59968
Katharia l'haru 1.625994 14.83906 0 180296 2.79368 0.047208 2.958091 8.45\679 \3.2\\23
Rana Tharu 11.52888 1.934922 2.591303 2.548486 0.636466 7.09765 38.11843 11.82609
Danguria(B) 10.99366 4.43276 6.2\5237 391.2288 3.571503 0.022738 14.02896 0 003288
Danguria 8.266575 27.19215 15.88918 303.331 1.718826 4.963945 19.79603 1.907203
Pachchimaha 2.08089 23.78175 I 1.34633 342.4747 0.752706 3.991829 2 990632 1.654925
Toto 36.74408 13.48364 0.703679 333.657 3.522\7\ 9.141742 27.98255 3.299776
MIZO 2.450938 12.40257 0.133762 5.726444 I. 782425 18.09615 0 235881 116.3543
Ho 5.961193 19.64705 0.987778 4 970542 4.683623 1.256438 34.1564 0.982124
Tipperah 5.467635 19.20867 15.96491 \0.23781 0.968436 4.739745 0.282451 2.503677
Badaga 33.32307 7.469853 3.093942 362.164 15.23141 25.77815 4.169706 3.469847
lru\a 22.5118 3.774505 18.80025 347.0716 16.34962 2.233618 3.253439 3.581462
Kota 20.77306 5.03591 2.310728 357.7866 13.7362 16.09358 22.36103 0.129357
Kuru mba 17.31585 2 922015 33.26448 393.767 6. 782337 0.127005 13.57406 1.899205
Toda 8.942353 81.23595 2.45691 339.7362 5.267575 4.96658 33.64218 22.42953
KondaReddi 2.533589 I 257963 13.38705 10.23781 3.424936 18.45381 1.772266 0.942476
Koya Dora 3.729544 0.039182 21.44862 1.443164 0.005577 4. 725637 5.989985 () 003288
x' (0.05) = 3.841 for 1 df (degree of freedom)
105
Tuhlc -1.17 Chi square \'a lues (I df) for the pair-wise comparison of Gene
frctJucncics of Bison-horn Muriu with twenty three populations of India -- ---- ~- - - - -
I PopuiHficm Alu no! Alu Alu Alu Alu Alu Alu Alu I
PLAT NliC Al'<l ACt: C04 I)J FXIIIH I'V 92 - ---~--~ ~~ -~ r------
lla!tm 11 06794 1.104961 0.202961 340.MY8 12.10666 5 524417 0.886588 0.294294
-- ~- ----
l\.mna1 5.57045 ·1.25695·1 o 36X2DJ 428.4(> 73 12.04168 1.685482 0.620432 2.01361 -~ --r----
( 'hinda Hbtmiia 1.]24663 21.5~307 6.009194 2Y8. 737Y 36.9248 I 1.790203 3.58322 0.5Y4524
( 'haukatia llhun.Jia 9.769535 5.682251 0.127801 316.411 0.308046 1.163711 3.802662 6 446YOI
---·
Mundu 14.75'!65 5631309 0 223883 317. 1()55 1.104063 6.268311 0.376462 1.076044
l.odha 3.375818 2Y.01994 15.79519 333.8028 0.133103 0.361894 0.07242 0219048
Santa! 4.303931 0.195187 6.211323 279.7094 0.286589 I 776116 0.571882 9.269972
Kathana l'haru 0.935267 6.248737 3.685697 0.532799 9.304959 O.Gl \605 11.43921 19.90721
Rana Tharu 9.751778 0.072683 0.940262 0.188888 15.09045 0.428834 45.70923 19.40177
Danguria (B) 9.269909 0.177747 0.017325 430.1534 1.892258 5.46654 18.7375 0.984125
Danguria 6.775331 15.35592 3.069129 341.0046 2.297674 0.205363 24.46488 4 949\58
Pachchimaha 1.246583 11.79424 0.945212 380.6781 4.589611 17.2076 4.970021 4.798494
Toto 34.53043 5.799664 2.403924 37\.9152 1.133729 1.528167 33.52134 1.03977
Mizo 1.654712 5.137745 3.424862 3.050847 1.84637 7.791183 0.788278 106.2427
llo 4.575134 8.403687 1.387494 2.646154 0.201498 8.724725 39.53685 0.063629
Tipperah 4.218149 9.021462 2.226254 5.472 16.40103 0.20289 \.028969 6 410969
Badaga 31.06794 I 304383 19.04676 400.6566 0.59194 53.55288 2.742874 1.062001
lrula 20.4046 13.51357 3.63659 385.3333 47.87018 13.30878 I. 978205 1.117602
Kota 18.7105 0.480593 0.900481 396.2903 42.76758 4.836583 19.60007 \.5080\4
Kurumba 15.30882 12.02745 II .26318 432.5838 31.46779 6.291 136 18.05277 5.399638
Toda 11.63767 61.24938 17.28644 377.8605 0.708979 0.059324 30.52776 16.07023
Knnda Reddi 1.585716 8.213897 1.332169 5.472 1.639327 42.47302 3.369248 0.00596
Koya Dora 2.593707 2.369963 5. 735894 0.000775 10.55351 0.008362 8.971801 0.984125
X' (0.05) = 3.841 for I df (degree of freedom)
106
T~thlc ~.II! ('hi squ11rc ntlucs (I df) for the pair-wise comparison of Gene
frl"IJUl'lll'ics of MuriH with twenty three popuhttions of India
Populnfion Alu mf Alu Alu Alu Alu Alu All• Alu
NIK' AI'O A('E Cl)4 I> I ~'X IIIII I'Y 92 I'LA:I'
I -- ---- - -- -- - ----
i J l.1lha 21 xww () 2·1').1)) 5 .1.10'i2 I 31-IOlXI> 2 62·CXll .l1N971X 0 70X'IJ I 1.02925
I ------- -- --- -------- ---- 1-------
1-.arll.tr 2 n.J(JIJ\ 2 062573 5 212·12-1 39tJ.Xfl(J6 2 .. 175X-I () 92225 0.474X45 .1.4571 X I
- ~----
< 'hl!lda Bhunpa 0 2MUXJ 16.2X4X7 20.92199 27-1.2454 211.05143 I. J3(, 7 13 3 5R740 I 1.371]51 --- -- --~---
( 'h,ur~a(r,I/Jhtm]ia 5.51734'! 3 27259 4.R576X2 2lJ0.956 !.] 15432 0.6162X3 3. 7R24R9 3.953739
- ---
Mnnda 10.0)245 4£>.490 12 3.22X949 291.]396 0.2141 97 4.799949 0.4219R7 0.34R338
----
l.odha I J 16292 22.24527 37.231X2 307.2855 1.512393 0.660809 0.041781 ()
--
S:mtal 2.09592 0.6'! 1525 0.023826 256.9131 0.879977 1.175177 0.469986 11.21459
1--Katharia 'J haru 0.080309 X.266645 0.493963 0.102716 1.737534 0.019747 10.37292 1498431
Rana r!wru 5.188644 0.069211 3.566203 0.545254 3.929458 0.805452 42.05654 13.72102
llanguna(R) 4.947525 0.877508 7.598008 401.5271 0.298758 3. 776439 16.75398 0.16674
Danguria 3.4213 18.14134 17.60958 314.3385 0.030006 0.459422 22.53974 2.774823
Pachchimaha 0.137179 14.52368 12.95141 352.8071 0.112974 13.54762 4.335012 2.527739
Toto 25.94436 7.642728 1.242044 344.1827 0.456701 2.07585 31.00695 2.111946
Mizo 0.422892 6.881821 0.397608 1.231793 0.063938 8.639467 0.650295 108 0808
llo 1.775318 10.92249 1.548396 1.067917 1.205427 6.807971 37.25144 0.462454
Tippcrah 1.756685 11.39306 17.8188 2.213943 4.640538 0.165533 0.82392 3.568933
Badaga 21.83039 2.588889 1.973807 372.4048 6.598931 44.73641 2. 738978 2.191767
lrula 13.33128 8.961808 20.71711 357.349 25.42998 10.25038 2.000273 2.272464
Kota 12.08622 1.352842 3.206308 368.0767 22.12081 5.66221 18.77559 0.461435
Kuru mba 9.233426 7.741216 35 52569 404.0578 13.66547 4.464701 16.20175 2.855379
Toda 1589051 66 55075 1.487468 350.0426 0.995395 0.248564 29.07822 18.43474
KondaRcdd1 0.228407 4.878318 15.13778 2.213943 0.322345 35.12982 2.880306 0.355708
Koya Dora 0.669223 0.874479 23.35167 1.362148 I. 736869 0.132025 7.883451 0 16674
X' (0.05) ~ 3.841 for I df (degree of freedom)
107
Tahl~ 4.1') AHfa)ll" h~t~rul.y)lusity and Standard ~rrur (SE} hased un dght Alu
in\l·rtiun/tldetion polymorphic loci in each of the twenty six populations of India
indutling pn•st·nt stud~· - .- ... - --- ---- --" ------
Tribe Avcrag~ hct~rozygosity + SE
·- - ----------- ---- -- ---- -· - ------
,\bujhm;u·ia 0.4~002~ 0.048362 - -- ·-· -· _, ____ . ---
Bison-horn Maria 0.381794 0.051725 --
Muria 0.398969 0.057396 .... --- --------
11alha 0.431461 0.038327 ----- --------
Kamar 0.417061 0.056279 -----
( "hinda Bhw1j ia 0.410993 0.52875 ..
( 'haukatia 13hunj ia 0.412454 0.038639 - --
Munda 0.407399 0.058939
Lodha 0.371705 0.060059
Santa! 0.409005 0.054646
Katharia Tharu 0.354340 0.056940
RanaTharu 0.355311 0.054665
Danguria (B) 0.388064 0.052086
Danguria 0.335785 0.050063
Pachchimaha 0.379227 0.054403
Toto 0.340662 0.056811
Mizo 0.305016 0.074116
Ho 0.357845 0.062897
Tipperah 0.358737 0.061036
Badaga 0.401724 0.056788
Irula 0.424503 0.051585
Kota 0.364882 0.061511
Kurumba 0.384225 0.059128
Toda 0.317344 0.063969
KondaReddi 0.411393 0.059770
Koya Dora 0.383468 0.052495
lUll
Tuhl~ .t.2tl R~'ult' of~~~~~ di\·~rsity analysis of tw~nty six populations of India
fur itHihiduallnci and fur ulllnci considered jointly
Lucu' II, 11, (;~I -- -- ---··
,\lunlt Nll( · 0.49l!461 .47l!l63 0.040721 - - -- - -- ------
--
, i\lui\1'0 0.40X:l39 0.161629 0.114191
I
I
-
_ .. .. - ·- f----·
i\ lu ;\( '1'. 0.454970 0.434025 0.406036 . __ , _______ _, .--·
-
i\lu ('J)4 0.479319 0.046493 - -- - ---------- f--
i\lu PI 0.471711 0.449778 - ... ·--·· -- -----·----"
;\lu J-'XIIlll 0.387439 0.365834 -------
i\lui'V92 0.489010 0.443972
i\lu PLAT 0.4 79663 0.435912
i\Jlloci 0.458614 0.376976
H 1 ~Total genomic diversity among the populations
H ~Diversity between individuals within populations '
Gsr ~Between population genomic diversity
0. 903002
0.046496
0.055764
0.092101
0.091213
0.178011
Table 4.22 References of Populations in India taken for comparison
S.No State Tribes Reference
I. Chhattisgarh Abujhmaria Present Study
2. Chhattisgarh Bison-hom Maria Present Study
3. Chhattisgarh Muria Present Study
4. Madhya Pradesh Halba Mukherjee et al., 2000
5. Madhya Pradesh Kamar MukheJjee eta!., 2000
6. Madhya Pradesh Chinda Bhuqjia Mukherjee eta!., 2000
7. Madhya Pradesh Chaukatia Bhunjia Mukherjee et al., 2000
8. West-Bengal Munda Majumder eta!., 1999
9. West-Bengal Lodha Majumder et al., 1999
I 0. West-Bengal Santa! Maj umder et a!., 1999
11. Uttar Pradesh Katharia Tharu Sharma & Shukla, 2004
12. Uttar Pradesh RanaTharu Sharma&. Shukla 2004
13. Uttar Pradesh Danguria(B) Sharma & Shukla 2004
14. Uttar Pradesh Danguria Sharma & Shukla 2004
--·
-
Table 4.21 G
enetic Distance o
f twenty six populations oflndia
S no 2
3 4
5 6
7 8
9 10
11 12
13 14
15 16
17 18
19 20
2'
22 L~
i~ t:
2 0 0038
3 0 002
0.0021
4 0 0684
0.0761 0.0807
5 0 0845
0.0926 0.0983
0.004
6. 0.0865
0 0972 0.1033
0.0121 0.008
7 0.0682
0 0739 0.08
0 0058 0 007
0.0135
8 01121
01233 0.1282 0.0172
00125 0.0154
0.0141
9 0 0966
0 0992 0 01104 0.0147 0 0091
0.01104 0 0101
0.0103
10 0 0854
0.0912 0 097
0 0071 0 004
0 0208 0.0118
0.0219 0.00191
11 00086 0.0095 00071
0.085 01051
0.1136 0.0862
0.1407 0.1191
0.106
12 00017 00134
0.011 0.0817
0.1025 0.1095
00864 0.1329
0.1122 01063
0.038
13 0 0876
0 0938 0 1008 0 0058
0.0042 0 0195
0.0097 0.019
0.0133 0.0062
0.0994 0.0963
14 0 0886
0 0905 0 0995
0.0132 0.01326
0.031 0 0178
0.039 0.0224
0 0156 0 09
0 0885 0.0051
15 0 0844
0 0904 0 0984
0.0097 0 0088
0 0221 0.0121
0.0299 0.0207
0.0108 0.0938
0 0967 0.0038
0.0055
16 0
10
41
01
07
4
01134 0.0128
00135 0.037
0.0231 00343
00273 00111
01094 0.1035
0.0068 00074
0.0152
17 00445
0.0357 00634
0.1249 01403
01512 0139
0.1895 0.1606
01269 0.0566
00587 0.1496
0.1462 0.1519
0.1464
18 0 0168
0 0148 0 0139
0 0997 0.1215
0.1367 0.1093
0.159 0.1374
01159 0.0107
0.0103 0.1121
0 1056 0.1099
0 1185 0 0429
19 00111
00077 00072 0094
01154 01177
0.0967 0.1526
0.1261 01173
00051 00095
0.1136 0.1033
01064 01238
00445 0.0132
20 00956 0
10
41
01
08
3 00122
00141 00322
00148 0.0235
0.0319 0.0087
0.1169 01206
00137 00279
00148 00231
01556 0127
01291
21 00851
00964 01011
00057 00059
0.0029 0.0113
0.0141 00135
0.0155 0.1092
0.1049 0.014
00257 00166
00275 01483
01279 0.1139
00206
22 01114
01195 01231
00165 00117
00155 00164
0.0256 0.0235
0.0191 0.1277
0.1301 0.0202
0.0259 0.0217
00261 01709
01603 01342
00273 00139
23 01094
01196 01262
00127 00085
00122 0.0178
00141 0.011
0.0206 0.1239
0.1154 0008
0.016 0.0136
0.019 01837
0.1414 0.1352
00294 00096
002
24 0 1069
0 1072 0 115
0.0458 0.0399
0.0595 0.0445
0 0809 0.0657
0 0287 0 122
01405 0.0439
0.0414 0 0332
0 0498 01353
01429 0 1293
0 0392 0 0574
0 0395 0 0726
25 00093
00101 001
00939 01141
0.1142 00964
01367 01209
0.1153 O.Q18
0.0184 0.1141
0.1174 0.1103
0.1344 0.0501
0.0128 00142 01195
01098 01512 01337
01487
26 0 0061
0 045 0 0062
0 0738 0 092
0.0915 0.0764
0 121 0.0957
0 0965 0 0086
0 0065 0.0921
0.0871 0.0904
0 1047 0 0435
0 0136 0 0061
0 1122 0 0905
0 1197 0 1088
0 1257 0 0094
s No show
s the popula\lons correspondmg to Table 4.13
~
110
' I' I :nar l'rad,;\h l'at:ht:himaha i Sharma & Shukla. 2004
Ill 'Jorlh hosl I olo C 'hakraharly .:1 al.. 2002
' - --- --~~ --------- r---17. I North Lasl Mim ( 'hakraharly .:1 al.. 2002
--
IX. Norlh 1-:asl flo Chakraharly cl al.. 2002 -·
' I
I') Norlh h1s1 'lippcrah Maj unHJcrel a I_:: 1999 I ---
~0. Soulh India Badaga Vishwanathan ct aL 2003 ---- ----·- --- ----
21. South India lmla Vishwanathan eta!.. 2003 ---~-- -- -- ·----~--- ·--·-------
00 Soulh India Kola Vishwanathan cl al.. 2003 --· n South India Kummba Vishwanathan et al.. 2003
- - --
24. South India Toda Vishwanathan cl al.. 2003 - -
25. South India KondaReddy Veerr~u et al.. 200 I
26. South India KoyaDora Veerr'\iu et al., 2001
4.8 Comparison with other Indian Populations
To determine the genetic relationship with other populations of!ndia, we compare our
results of eight Alu insertion/deletion polymorphic loci with available twenty six populations
oflndia. These loci are common with our studied populations & populations considered for
comparison.
Brief descrition of26 populations taken for comparison is given in Table 4.12. Locus
specific allele frequencies are presented in Table 4.13. Table 4.14 shows the number of
individuals studied at each locus examined for comparsion. Table 4.15 shows the
heterozygosity at individual loci based on Alu insertion/deletion polymorphic loci in 26
populations of India. Among them six tribes from Central India (M.P. & C.G.) (including
present study) belong to Dravidian and one tribe to Indo-Aryan language families, five
population from eastern India (W.B. & Orissa) belong to Austro-Asiatic language families,
two populations from N-E region (one belongs to Austro-Asiatic and one belongs to Tibeto
- Burman language family) and seven populations from South India belong to Dravidian
language family).
The allele frequencies of Alu mt NUC among Indian populations varied from a low of
0.204 in Toda of south India to as high as 0.625 in Badaga of south-India and Halba of
Madhya Pradesh. The allele frequency of Alu mt NUC of the present study is within the
range of distribution oflndian population.
The Alu APO allele frequency among Indian populations varied from a low of0.300 in
Munda to as high as 1.0 in Toda of South India. Our findings are within this range. Alu ACE
Ill
,tilde ""'l"cnc:~ am on~ Indian populatitlllS varied from a low of0.-\(>0 in Bada)!a of South
lndta 111 "' hi)!h as IJ.X59 111 l.odha of W.JI.<Jur lindings an: within the range of Indian population
Rc"dl;, of/\lul'D-1 polymorphism in population studied in India varied f(mn a low of
IJ in Kola & Kununba of south-India to as high as I in Toto. Mizo. llo and Konda Reddi.Our
results ranges bet ween them.
'I he /\lu D I allele fre<Jucncy in the Indian populations varied from 0.230 in Badaga to
IJJt20 in ( 'hinda Bhunjia. Our study gave a range of0.2627 in BHR & 0.4057 in ABM, thus
within the range of Indian populations.
Alu FXIIII3 allele frc<juency in the Indian populations varied from as low as 0.461 in
lladaga to as high as 0.944 in Toto. The findings of the present study ranges between them.
The results of Alu PV92 polymorphisms in available Indian populations varied from as
low as 0.255 in Toda to as high as 0.933 in Toto. The allele frequencies of populations
under study ranges between them. Likewise allele frequency of Alu PLAT in the available
Indian populations varied from as low as 0.033 in Toto to as high as 0.814 in Katharia
Tharu.
Tables 4.16, 4.17 and 4.18 show the x' values for the pair-wise comparison of gene
frequencies of Abujhmaria, Bison-horn Maria & Muria respectively with other populations
of India.
Of the total 4 72 pair-wise comparison of gene frq uencies of the populations under
study with twenty three populations oflndia, 247 (52.33%) are statistically non-significant.
Most of the non-significant comparisons are between Muria and other Indian populations
taken for comparison.94 of the 184 (51.63%) non-significant differences are between these
groups. The next population that showed non-significant difference is Bison-horn Maria.
BHR showed non-significant difference in 79 of the 184 (42.93%) comparisons. Abujhmaria
accounted for 74 out of 184 (40.22%) non-significant comparisons.
Abujhmaria significantly differs with Halba, Kamar, Danguria(B), Tipperah, Irula and
Kurumba at 5 loci (62.5%), with Chinda Bhunjia, Lodha, Danguria, Toto, Ho, Badaga &
Kota at 6loci (75%), with Chaukatia Bhunjia, Munda, Rana Tharu, Pachchimaha & Mizo at
4loci (50%), with Santa!, Katharia Tharu, Konda Reddy & Koya Dora at 3 loci (37.5%)
and only with Toda at seven loci (87.5%). Alu CD4locus contributed for most of the significant
comparisons (20 out of23 i.e. 86.9%). Abujhmaria shows greater similarity with Santa!.
Katharia Tharu, Konda Reddy & Koya Dora.
Bison-horn Maria differs with Halba, Kamar, Chinda Bhunjia. Chaukatia Bhunjia.
Munda, Santa], Katharia Tharu. Rana Tharu. Danguria (B). Toto. Mizo and llo at -\loci
112
(:'O"ul. \\ith I mlha. Konda R..:ddy & Koya Dora at 3 loci (37.5"1.,). with Danguria.
J'arhdlimaha. I ipprrah. lladaga. lruht & Kola at 5 loci (62.5%) and with Kuru mba & Toda
.11 l> lo~:i ( 75'!io).Aiu APO and Alu CD41oci contrihul<:d for most ofsigni!lcant comparisons
11 H out of::'J i.e . .1'1.11'%). Bison-horn Maria shows greater similarity with J.odhu. Kondu
Rrdd) & Koya Dora.
Muria differs with llalha.l'achchimaha. Kola & Todu utSioci (62.5%), with Chinda
Jlhunjia. Chaukatia Bhunjia, Munda, Rana Tharu, Danguria (B), Danguria. Toto & Badaga
at4 loci (50'X,), with Kumar, Suntul & Koya Dora at 2 loci (25%) with Lodha, Katharia
J'haru. Mizo, I lo, Tippcruh & Konda Reddy at 3 loci (37.5%) and only with Kurumba at I
locus (87.5%). Alu CD4locus contributed for most of the significant comparisons (16 out of
2J i.e. 69.56'Yo). Muria shows greater similarity with Kamar, Santa! & Koya Dora.
Locus-wise estimates of heterozygosity of available insertion/deletion polymorphic loci
among Indian populations are presented in Table 4.15. A persual of table reveals that most
populations show high level of heterozygosity for Alu mt NUC while lowest for Alu CD4. The
populations exhibited maximum variation for Alu APO and on the other hand Alu mtNUC even
though exhibited highest heterozygosity showed lowest difference among different
populations. There was great intra-locus variation in values over loci in the present study as
well as in other reported studies from India. Alu CD4 was the least heterozygous locus and
Alu mt NUC the most heterozygous locus in all populations.
The average heterozygosity (H) with their standard errors are presented in Table 4.19.
It was found least in Mizo (0.305) & most in Halba (0.43). In the remaining populations it
varied between them.
The populations under study showed the estimates of average heterozygosity ranges
between 0.3817 in BHR to 0.4200 in ABM. The H value is MUR is 0.3989. This shows that
the ABM are comparatively more heterozygous than those BHR & MUR .
Estimates of various measures of gene diversity analysis viz., Hr. H, & G5r among the 26
populations of India (including present study) are presented in Table 4.20.
In the present material, intra population gene diversity (Hs) varied greatly from a minimum
of 0.0464 at Alu CD4 to a high of 0.4 781 at Alu mt NUC locus with a mean of 0.3 769 over all
eight loci. Inter-population gene diversity varied from as low of 0.3874 at Alu FXIIIB to a
high of 0.4984 again at Alu mt NUC locus, with a mean of 0.4586 over all eight loci.
Co-efficient of gene differentiation (Gsrl also varied greatly over loci in the Indian
populations. Alu mt NUC locus being the least differentiated locus (0.0407) and Alu CD4
being the most differentiated locus (0.9030), the remaining six shm,ed the moderate values of
IIJ
t 1 u.w~l..' u ().l(,.l t~l u -HI61 I h~.: a\ cra!!,C mca,ulc tlt' )!.CI\C:tlc..: dtllcn.:ntiation O\ cr all tin: Cl)!.hl
\1'\.' \0 I 7XO) \\a' luund tu he In''· \\hll:h imhcatc~ lln\ cvolutlmHll") dtvl!rgcm:...: anHHlg the
1'a1n' ,, .... gcnctu.: dl' .. tarn.:c hct\\CCII the .'!llHiy populations as well as :!6 tribal populations
nl lndta \\L'rc calculated from the allele frcqucncic.'l using the D, distance measure (Nci ct a!.,
I '172) arc!-!'""" 111 Ia hie 4.21.11 may he noted !hal Abujhmaria shows grcalcsl gcnclic dislance
"''" Munda (0.1121) '"compared lo olhcr populations. The genetic distance is also higher
bclwccn 1\BM a11d Kola (0.1\14). 1\BM and Kurumba (0.1094). 1\BM and Toda (0.\069) &
i\BM and 'loto (0.\041) respectivcly.This indicates that the gene diiTcrenccs between i\BM
and lhesc popu\;llions arc high.Cicographieally also these populations are distant from A 13M.
The lowest gc11ctic distance is observed between ABM and MUR (0.002). which indicates
!hal the gene diiTerenccs between ;\13M and MUR arc very small accounting for not more
!han 0.2 percent of total genetic variation. ABM also shows lower genetic distance with BIIR
(0.0038) and Ran a Tharu (0.00 17).Gcographically also ABM. BHR & MUR are close to each
other & belong to major Gond tribe.
It is marked that BHR shows max1mum genetic distance with Munda (0.1233) as
compared to other populations, but the distance between BHR and MUR is very small, indicating
almost negligible gene differences between two populations accounting for not more than 0.21
percent of total genetic variation. BHR also shows considerably higher genetic distance with
Kota (0.1195) and Kurumba (0.1196), which indicates that the genetic differences between
BHR & Kota and BHR & Kurumba are higher. It is marked that besides MUR, BHR also
shows lower genetic distance with Tipperah (0.0077) and Katharia Tharu (0.0095), which
indicates that the genetic differences between BHR & Tipperah and BHR & Katharia Tharu
are smaller.
The highest genetic distance is observed between MUR and Munda (0.1282), which
indicates that the gene differences between MUR and Munda are very high. MUR also exhibit
higher genetic distance with Kurumba (0.1262) and Kota (0. !231 ). It is marked that MUR
shows least genetic distance with Koya Dora indicating almost negligible gene differences
between two populations accounting for more than 0.62 percent of total genetic variation.
MUR also exhibits smaller genetic distance with Katharia Tbaru (0.0071) and Tipperah (0.0072),
which indicates that the gene differences between MUR and these populations are smaller. The
three tribes under study (Dravidian) show the highest genetic distances with Munda. which
belongs to the Austro-Asiatic language family.
114
I" ICAmdr JL S<~ntd I L Hlllhll
Chaukalld811UIIj id
-Toto - D11nguria
D11nguriall rt~chc!llma Ia
'----- Dt~daya
....--- J1unda '---Lodha
J(urunha r-----= ChI ndi\Btmnj ia ~Irult\
L----Rola L---------------------Toda
.--------------------11izo ~__ _____________ L ---Ho
____________ _j1----- J<ouddRedtl 1
-·C: Ita thar i Tharu llana Tl1aru li p}lerah l<ot)dDor"
r B ison-hor-nl1dr ia Abujltmt\r ia J1ur ia
Figure 4.12 Neighbour-joining tree depicting genomic affinities among the 26
tribal populations in India based on eight insertion/deletion polymorphisms
An unrooted neighbor-joining tree was constructed from this distance matrix (Figure
4.12). It is pertinent to note that three tribal populations of the present study viz. Abujhmaria,
Bison-horn Maria, Muria and Kamar Konda Reddi, KoyaDora, Badaga, lrula, Kota, Kurumba
& Toda are linguistically belong to Dravidian language family, while Halba, Chinda Bhunjia,
Chauktia Bhunjia, KhatriaTharu RanaTharu, Danguria (B), Danguria Pachchimaha belong to
Indo-European language Family. Munda, Lodha, San tal Toto, Mizo and Hoare belong to Austro
Asiatic linguistic family and only one tribe Tipperah belong to Tibeto-Burman language family.
It is evident from Figure 4.12 that dendrogram consists of two main clusters A and B.The
cluster A is further sub-divided into two sub-groups, One group consist of Kamar and other
consists of Santa!, Halba, Chaukatia Bhunjia, Toto, Danguria, Danguria (B), Pachchimaha,
Badaga, Munda, Lodha, Kurumba, Chinda Bhunjia, lrula & Kota. Cluster B is also sub-divided
into two sub-groups. One group consists of of Ho, Konda Reddi, KathariaTharu and
RanaTharu, Tipperah, Koya Dora, Bison-horn Maria, Abujhmaria and Muria.The Toda & Mizo
positioned as sub-groups in cluster A & B. Abujhmaria, Bison-Horn Maria and Muria the sub
groups of major Gond tribe & are linguistically similar and live in close geographical proximity.
which is also reflected in the dendrogram, as they positioned closer to each other. It is interesting
to note that the Koya Dora, the Dravidian speaking tribe of South India is closer to lhc
populations under study, who also belongs to Dravidian language family & hislor~ also revcab
their migration from South India to Central india (Dubey.2000). Ho\\evcr. the other Dra\ idian
'PL-.1h.1n~ -.,pcah..ln).! '-lollth lndJan tnhc' arc di,tantl) placed in lkndrogram.Surpli'>ingly l\H\
Jnd11 .11~.111 "P"-'<I~lllg pnpulalll)/1'> oll!llar Pradc..,h.onc I iht:tn~Burman -,pca~ing tribe of North
1 ,hl lnd1a & lHlL' Athlnl-A..,I<IIIL' -.,pca~ing trihc nf cast India arc clu.,tcrcd \\ith the present
... tud~ I hu-.,, th1.., allaly..,j., t'l'\caJ., that the IJra\idian and Au.,tro~A~iatic :-.pcal..ing, trihal:-. arc
,L!,L'IIL'Iicall~ l..'lo.,L·r thaulndo Ar)an & Tihcto Burman "'ipcahing trihc"i.
l hi:-. rc-.,tdh indicate that in spite of their geographical proximity, the tribes remain isolated
111 n1o..,t of the region:-. with reference to historical. socio-cultural and genetic prespcctives.
/\gain the trihc> living in different regions arc not genetically similar. Their genetic differentiation
i> inf'lucnccd more by geographical proximity than by other f'ttctors. The tribes who arc closed
geographically have smaller genetic distance than those living apart. For example the tribes of
the present study (Abujhmaria, Bison-horn Maria and Muria) and South Indian tribe (Koya
!lora) arc genetically closer than the tribes in Uttar Pradesh (Katharia Tharu and Rana Tharu),
Tripura (Tippcrah).Thc genetic distance between tribes is correlated more with geographic
continuity than with linguistic affinity implying that exchange of genes occurs more often
between neighbouring tribes than between tribes of same language groups living at a distance.
The overall inference from the dendogram indicates that the populations of India either
from eastern, western, central India and southern India, geographical proximity as well as
languages has stronger effect than socio-cultural proximity in determining genetic affinity.
Table 4.23 Allele frequencies at five insertion/deletion polymorphic loci in
seventy one World Populations taken for comparison
Population Alu Alu Alu Alu Alu
ACE FXIIIB PLAT Dl PV92
Mbuti 0.320 0.030 0.240 0.590 0.350
African Pygmies 0.221 O.oJ5 0.221 0.338 0.309
Afro Caribbeans 0.524 0.310 0.286 0.405 0.143 I
Berber 0.412 0.344 0.526 0.233 0.269
Moroccans 0.292 0.236 0.547 0.396 0.274
Arabs 0.290 0.310 0.390 0.370 0.270
Turks 0.441 0.473 0.427 0.284 0.256
Pakistani 0.440 0.230 0.510 0.170 0.300
Shrilankan 0.623 0.613 0.598 0.264 0.481
Moor, Shrilankan 0.591 0.625 0.642 0.265 0.490
Han 0.670 0.710 0.350 0.170 0.860 ---~- ,_
Sibo 0.583 0.825 0.533 0.283 i O.l\50 i
- -- -- -
116
l\.lltl..',ll\ 0.500 0.!<46 0.421 0.15-l 0.846
' I \ghw· 0.650 0.776 0.529 (). 2 ()() 0.54 7
' I {ai\\an\..'~ 0.500 0.970 0.640 <UKO 0.900
·-- ·- - - _,_ __ - ·-
I ilipino 0.510 0. 720 0.630 0.360 0.800 - --------- .-- r------ ---- --- ·- ,_
Moluccas 0.670 0.7!<0 0.560 0.190 0.690 -- ._ __ ,
N usa ·r l:ngarras 0.640 0.810 0.380 0.190 0.500 ----
.lav;_mes~.: 0.&60 0.920 0.390 0.420 0.840 ... _____ , __ ------
Malaysim1s 0.640 0.730 0.500 0.270 0.720 --·--- ___ ,
Swiss 0.372 0.477 0.453 0.337 0.200 -·
llllsquc 0.369 0.510 0.524 0.476 0.194 ---
French 0.480 0.420 0.560 0.460 0.230
Greeks 0.260 0.500 0.552 0.409 0.190
Bretons 0.478 0.400 0.556 0.389 0.267
English 0.436 0.489 0.458 0.477 0.283
Abujhmaria 0.546 0.710 0.641 0.405 0.539
Bison-hom Maria 0.666 0.796 0.600 0.262 0.517
Muria 0.530 0.785 0.625 0.346 0.520
Halba 0.646 0.427 0.574 0.427 0.563
Kamar 0.640 0.420 0.535 0.420 0.554
ChindaBhunjia 0.796 0.620 0.556 0.620 0.407
Chauktia Bhunjia 0.649 0.289 0.729 0.289 0.412
Katharia Tharu 0.569 0.417 0.819 0.417 0.694
RanaTharu 0.623 0.443 0.792 0.443 0.821
Toto. 0.603 0.333 0.033 0.333 0.567
Mizo 0.731 0.286 0.586 0.286 0.862
Ho 0.460 0.452 0.713 0.452 0.565
Badaga 0.750 0.230 0.551 0.230 0.436
Irula 0.622 0.600 0.550 0.600 0.449
Kota 0.806 0.589 0.659 0.589 0.300
Kurumba 0.469 0.528 0.704 0.528 0.71J -----
Agahria 0.417 0.417 0.587 0.417 o .. ns I ~ -- --- - ~
117
I l.tt"dl OH!~ ' 0.6~5 0.4!(~ 0.645 OA6X I . - ·- I - - .
< 'halllar I 0 700 0. 500 0.411 0.500 0.5~0
< iaud 0.600 0.200 0.4~~ 0.200 o.:n:1
I odha 0.!!5'! 0.2X I 0.625 0.281 0.532 -- - -- --------- -
Malu,hiya 0.559 0.588 0.4!(5 0.588 0.515 --- -----
' I Munda 0.640 0.320 0.660 0.320 0.4X I
---- - --·~--- C----
San<al 0.521 0.292 0.417 0.292 0.563 ----- -- ---- - -- --- -------- ---1---
hUJti 0.433 0.406 0.718 0.406 0.656
'lippcrah 0.590 0.313 0.549 0.313 0.811 -
lycngcr 0.540 0.461 0.559 0.461 0.461 -----
lycr 0.570 0.420 0.510 0.420 0.430
V<UJniyar 0.620 0.410 0.570 0.410 0.490
Ambalakaran 0.480 0.370 0.650 0.370 0.390
Vcrrakodivellalar 0.663 0.288 0.605 0.288 0.488
Mvskoke 0.700 0.790 0.490 0.460 0.570
Alaskan natives 0.583 0.917 0.298 0.415 0.619
French Acadians 0.510 0.480 0.430 0.420 0.18
Inuit, Greenland 0.548 0.786 0.333 0.452 0.607
HispannicAmerican 0.545 0.705 0.625 0.364 0.523
Maya, Yucatan 0.680 0.900 0.650 0.450 0.790
Tuva 0.737 0.803 0.462 0.397 0.590
Yakut 0.578 0.836 0.524 0.551 0.698
Bury at 0.734 0.851 0.521 0.287 0.511
Papua New Guinean, Costal 0.660 0.300 0.160 0.170 0.360
Papua New Guinean, Highlanders 0.740 0.300 0.160 0.010 0.240
Australian 0.910 0.650 0.130 0.040 0.150
Guarani 0.829 0.935 0.710 0.394 0.783
Kaingang 0.543 0.872 0.675 0.706 0.793
I
I
liM
l'ahlc ~.H The numhcr of indi,·idual' at each locus 'tudied in sc\·cnty one World
l'npulatinn' l'\llmint•d fur t·nmparison
l'npulatinn Alu Alu Alu Alu Alu
A('E ~·xum I'LAT Ill I'V'H ---- ---
n n n n n - - -- -
Mhuli 14 .14 14 14 .14 - - - -- -- - --
Ali·icanl'ygmics 6S 6S 6S 6S 68 - - -- ----- f------
;\ liu ( 'ari hhcans S4 84 84 84 84 - - ---~--- ---- --- - f----
1krhcr so so 80 so 80 -- -- - -- ---- --
Moroccans 106 106 106 106 106
Arabs 100 100 100 100 100 ---- --------
Turks 86 86 86 86 86 -------
Pakislani 84 84 84 84 84
Shrilankan 96 96 96 96 96
Moor. Shrilankan 102 102 102 102 102
Han 98 98 98 98 98
Sibo 120 120 170 120 120
Korean 26 26 26 26 26
Uyghur 170 170 170 170 170
Taiwanese 92 92 92 92 92
Filipino 94 94 94 94 94
Moluccas 96 96 96 96 96
Nusa Tengarras 180 180 180 180 180
Javanese 64 64 64 64 64
Malaysians 94 94 94 94 94
Swiss 86 86 86 86 86
Basque 206 206 206 206 206
French 106 106 106 106 I 06
Greeks 102 102 102 102 I 02
Bretons 90 90 90 90 90
English 188 180 192 176 184 -- ,J
11'1
\butllln,U1.l i S-1 50 S.l SJ S I
i!J,on-homl\ !aria 60 59 60 59 sx I \ h1ria 4<J 49 4X 49 49 I
llalha 4X 4X 47 4X 4X --- ---
Kamar 57 57 57 57 57 1- -- - ---- --
( 'hinda Blumjia 27 27 27 27 27 - --··---- -· -------------~
( 'hauktia Bhlmjia 37 37 35 37 37 --
Katharia Thm·u 36 36 36 36 36 --- ----- -- -·---r-·
Rana 'l'haru 53 53 53 53 53 - ---- --~---
'!(>to. 29 30 30 30 30 ---
Mim 54 29 29 29 29
llo 50 54 54 54 54
Badaga 50 50 49 50 50
Irula 45 50 50 50 50
Kota 54 45 44 45 45
Kurumba 49 54 54 54 54
Agahria 24 24 23 24 24
Bagdi 31 31 31 31 31
Chamar 25 25 23 25 25
Gaud 15 15 15 15 15
lodha 32 32 32 32 32
Mahashiya 34 34 33 34 34
Munda 25 25 25 25 25
Santa! 24 24 24 24 24
I anti 15 15 16 15 15
Tipperah 39 39 41 39 39
Iyenger 50 50 51 50 50
Iyer 50 50 50 50 50
Vanniyar 50 50 50 50 50
Ambalakaran 50 50 50 50 50
Verrakodivellalar 43 43 43 43 4], ----- __ j
120
''"'"'" ! 100 100 100 100 100
·\l;"k.Ulll<l!ll L'' X4 !!4 H4 84 !!4 I I I· r,•nd1 Al."adiam 92 92 92 92 92
- - -
Inuit. ( ircenland X4 84 H4 84 84 -- --- --~- -· ----
-- -
I lispannic Amcricm1 XX XX 88 8X 88 - ---- - ---·---- --- +-- -· --f-------
Maya. Yucai<Ul 102 102 102 102 102 - ---- --------------- -----
l'uva 76 76 76 76 76 ------
Yakut 128 128 128 118 126 - ------------ - ---------1-----
lluryat 92 92 92 92 92 ---
Papua New Guinean, Coastal 96 96 96 96 96 -------
Papua New Guinean,llighlandcrs 136 136 136 136 136
Australian 138 138 138 138 138
Guarani 70 62 62 66 60
Kaingang 92 86 80 92 82
n =number of individual
Table 4.25 Heterozygosity at individual locus of five Alu insertion/deletion
polymorphic loci in seventy one World Populations
Tribe Alu Alu Alu Alu Alu
ACE D1 FXIIIB PV92 PLAT
Mbuti 0.4352 0.4838 0.0582 0.4550 0.3648
Afiican Pygmies 0.3443 0.4475 0.0295 0.4270 0.3443
Afro Carib beans 0.4988 0.4819 0.4278 0.2451 0.4084
Berber 0.4845 0.3574 0.4513 0.3932 0.4986
Moroccans 0.4134 0.4783 0.3606 0.3978 0.4955
Arabs 0.4118 0.4662 0.4278 0.3942 0.4758
Turks 0.4930 0.4066 0.4985 0.3809 0.4893
Pakistani 0.4928 0.2822 0.3542 0.4200 0.4998
Shrilankan 0.4697 0.3886 0.4744 0.4992 0.4807
Moor, Shrilankan 0.4834 0.3895 0.4687 0.4998 0.4596
Han 0.4422 0.2822 0.4118 0.2408 0.4550 I
I ----1
Sibo 0.4862 0.4058 0.2887 0.2550 0.497R -- -
122
ltt~dt U.-12-17 11.4579 0.457'1 0.4'179 0.4994 ' I
0..1848 l 'h;uttaJ 0.-1200 0.5000 0.5000 0.4968 -
! ( i.tud 0 .4KOO 0.32011 OJ200 0.4442 0.491 0
- -
I <~dha <J.24n 0.4040 0.4040 0.4979 0.4687 -- ------ -
~ lalu,hiya 0.49:10 0.4845 0.4X45 0.4995 0.4995 ~ - -- --
Munda 0.460K 0.4352 0.4352 0.4992 0.448K -~ -- -- ---------
Santa I 0.499 I 0.4134 0.4134 0.4920 0.4862 ----
hu1ti 0.4910 0.4823 0.4K23 0.4513 0.4049 ------ --- ·--- - -
!ippcnJ1 0.4838 0.4300 0.4300 0.3065 0.4951 --------
lycngcr 0.4968 0.4969 0.4969 0.4969 0.4930 -
lycr 0.4902 0.4872 0.4872 0.4902 0.4998
Vmmiyar 0.4712 0.4838 0.4838 0.4998 0.4902
Ambalakaran 0.4992 0.4662 0.4662 0.4758 0.4550
Verrakodivellalar 0.4468 0.4101 0.4101 0.4997 0.4779
Mvskoke 0.4200 0.4968 0.3318 0.4902 0.4998
Alaskan natives 0.4862 0.4855 0.1522 0.4716 0.4183
FrenchAcadians 0.4998 0.4872 0.4992 0.2952 0.4902
Inuit, Greenland 0.4953 0.4953 0.3364 0.4771 0.4442
HispannicAmerican 0.4959 0.4630 0.4159 0.4989 0.4687
Maya, Yucatan 0.4352 0.4950 0.1800 0.3318 0.4550
Tuva 0.3876 0.4787 0.3163 0.4838 0.4971
Yakut 0.4878 0.4947 0.2742 0.4215 0.4988
Btuyat 0.3904 0.4092 0.2535 0.4997 0.4991
Papua New Guinean, Costal 0.4488 0.2822 0.4200 0.4608 0.4608
Papua NewGuinean, Highlanders 0.3848 0.0198 0.4200 0.3648 0.4823
Australian 0.1638 0.0768 0.4550 0.2550 0.2688
Guarani 0.2835 0.4775 0.1215 0.3398 0.2688
Kaingang 0.4963 0.4151 0.2232 0.3283 0.2262
123
'luhl•· ~.2(• A'~ra~~ h~tcro1.y~osili~s and Standard error (SEl based on fin: Alu
inwrtiunilld<·liun pt>l)morphic loci in sc,·cnty one World l'opulations
l'upu Ia tiun An rage hclcrozygosil)' + SE -
' ' 0.07K979 \1huli 0.)1>4764
-· --
Ali·icanl'ygmi.:s 0 .. 120907 0.075lW7 ---- -- ---
/\fro l 'arihheans 0.4 I 4X I 9 0.045299 - - ------·-- ---------
Berber 0.439786 0.027 I 02 - -- --- -~ ------- --
Moroccans 0.43 I 210 0.025378 --- ------
Arabs 0.437347 0.015719
rurks 0.456361 0.024979 ~ ---
l'akisumi 0.4!2254 0.041761 - ---
Shrilankan 0.464998 0.019261 1------
Moor, Shrilankan 0.462509 0.019031
Han 0.368279 0.044112
Sibo 0.388218 0.049918
Korean .360910 0.058349
Uyghur 0.424558 0.037695
Taiwanese 0.335865 0.090559
Filipino 0.431978 0.031575
Moluccas 0.404869 0.033981
Nusa Tengarras 0.410661 0.042138
Javanese 0.326511 0.067949
Malaysians 0.432782 0.021479
Swiss 0.448330 0.033047
Basque 0.456288 0.036287
French 0.468249 0.028167
Greeks 0.436265 0.038147
Bretons 0.470522 0.019715
English 0.479866 0.018288
Abujhmaria 0.473859 0.015962
Bison-hom Maria 0.430691 O.OJ2JOR I
I --- J
124
\hu·r.t 0.-I~~K~~ 0 .ti:Hl2YI
llalha 0.-IXX~-1.1 0.006611
Ka111<tr 0.-IK% 79 0.00652:1 I
i ( 'hi11da Jlhunj ia 0.4S71X6 0.031 X 59
( 'hauJ..tia 1\lmnjia 0.417402 0.0 \6X65 - ----------.
Kat haria Th;mr 0.44297X 0.037673 ------ ----- ---------- -~---·---· ---
Rana 'l'haru 0.419989 0.043593 - ~-
"1<>10. 0.393419 0.082815 --- -- ·-- -- ----- ·-------
Mi/0 0.410827 0.045398 - ---- -·
llo 0.461246 0.023106 --·- -- ------·
Badaga 0.442810 0.034723 --
lnila 0.469656 0.022965
Kola 0.466807 0.012319
Kurumba 0.431116 0.034733
Agahria 0.497469 0.001149
Bagdi 0.475279 0.014306
Chamar 0.490311 0.015665
Gaud 0.425223 0.039276
lodha 0.409823 0.044966
Mahashiya 0.499625 0.003444
Munda 0.465159 0.012102
San tal 0.470675 0.019871
Tanti 0.478156 0.016579
Tipperah 0.434646 0.033854
Iyenger 0.501134 0.000804
lyer 0.495879 0.002343
Vanniyar 0.490667 0.004719
Ambalakaran 0.477253 0.007523
Verrakodivellalar 0.454231 0.018154 i Mvskoke 0.449970 O.O:P668 ---1
Alaskan natives 0.405225 0.64251 -----
125
ill" Ill h. \c·:lliJ:Uh 0.-l:ii>XOl (). ().j()() 7-1
/runt.! iJL"<"IIiaml I 0.-l:i~ liJt> 0.0.1000-1
' I I 0.0 1503'1 I llhpan!llc AnH:ricwt 0.-1711'17 i
\lay a. Yucatan lUll I ~I>'! 0.056'144
l111a 0.415611 0.015190 .
Yakut 0.4.17202 0.042885 ----- -- -~-~ ..
llwyat 0.4!2688 0.0454.16 . ~- ---- ---- ---- ---- -----
Papua N~w ( iuincan. ( 'ostal 0.378089 0.04!882 -~~ -- - --- -- -·
Papua N~w (iuinc;m,llighlandcrs 0.2927!6 0.072699 .. - ----- --~-~- -------
i\ustmli.u1 0.236216 0.063067 ~. ~--~----
<Juanmi 0.3294!9 0.06!351 -------
Kaingang 0.382555 0.047933
Table 4.27 Results of gene diversity analysis of seventy one World population
for individual loci and for all loci considered jointly
Locus H,.
A1uACE 0.487197
AluDJ 0.465362
A1uFXIIIB 0.497458
AluPV92 0.499906
A1uPLAT 0.499583
All1oci 0.489901
HT =Total genomic diversity among the populations
H, ~ Diversity between individuals within populations
G,T ~Between population genomic diversity
Hs Gsr
0.446437 0.083664
0.428752 0.078669
0.390324 0.215363
0.417164 0.165515
0.451903 0.095438
0.426916 0.128567
Table 4.29 References of Studied Populations taken for comparison
S.No Location Population Reference
1. Africa Mbuti Bowcock eta!., 1987
2. Africa African Pygmies Stoneking eta!., 1997
3. Africa Afro Caribbeans Stoneking et al.. 1997
4. Africa Berber Stoneking et al.. 1997
5. Africa Moroccans Stoneking ct al.. 1997 -- ---- - '
12b
I> -\_...i.l Arab' ( 'hbd ct a!.. ~00.1
' 7 A:-.ta Juri., ( 'oma' ~I al.. ~004
s 1\:-.ia Pakistani Ston~king ~~ ai.. 19'!7 i
!) Asia Shril.uli-:m Antun~t.-de-Mayolo ~~a!.. 2002
Iii Asia Moor. Shrilankan Antun~z-de-Mayoio cl al.. 2002
12 J·:ast Asia I i<m Mellon ct al.. I 995 ---- --- ------ . --
11 !·:astAsia Sibo Xiao cl al., 2002 - - ----- ---------
14 l ·:tL~t Asiu Korean Antuncz-dc-Mayolo ct al.. 2002 ---- - - --- - --·-
15 l'ast Asia llygbur Xiao ct al., 2002 --- ---- ---- --
16 !·:astAsia Taiwanese Melton et al., 1995 - '-----
17 East Asia Filipino Melton et al., 1995 - --
18 East Asia Moluccans Perna et al., 1992 ---
19 East Asia N usa Tengarras Perna et al., 1992
20 East Asia Javanese Melton et al., 1995
21 East Asia Malaysians Melton eta!., 1995
22 Europe Swiss Batzer eta!., 1996
23 Europe Basque Antunez-de-Mayolo eta!., 2002
24 Europe French Monson eta!., 1995
25 Europe Greeks Comas eta!., 2005
27 Europe Bretons Monson eta!., 1995
28 Europe English Mastana et al., 2003
29 Indian Agahria M~umder eta!., 1999
30 Indian Bagdi Majumder eta!., 1999
31 Indian Chamar Majumder et al., 1999
32 Indian Gaud Majumder eta!., 1999
34 Indian Tanti Majumder eta!., 1999
35 Indian Mahashiya Majumder eta!., 1999
38 NothAmerica Mvskoke Weiss et al., 1993
39 NothAmerica Alaskan natives Batzer et a!., 1996
40 NothAmerica FrenchAcadians Batzer et al.. 1996
41 NothAmerica Inuit Greenland Batzer eta\.. 1996 ------'
127
·L' '-'othAmcri<:a I lli~txumi<: .1\m~ri<:<m Batter <:tal .. 1 1l1l6 ---- --- -- . ----
-
.J' I ~othAmcri<:a 1\·laya. Yu<:atan Weiss <:tal.. liJIJ1
! .j.l SilX'ria lu\'a .i\ntunc/·tk-Maynlo ct al.. 2002
4~ SilX'ria Yakut Khitrinskaya d al.. 2001 - -
4fl Siocria llwyal .1\nluncz-de-Mayolo cl al.. 2002 - -- ---
--
41J ( kcania Papua New Perna ct al.. 1992
(iuincan. Costal ---- - --- - ---- -~----- ------
)II Oceania Papua New Guinean, Perna ct al., 1992
llighlandcrs - ------
51 Oceania Aust.rdlian Perna et al., 1992
52 South America Guarani Battilana et al., 2002
5:1 South America Kaingang Battilana et al.. 2002
4. 9 Comparison with World Populations
To determine the genetic relationships of the three ethnic populations of Bastar,
Chhattisgarh of India with populations of other regions of the World we have compared the
available data on five Alu insertion loci that are common with our study. These loci are Alu
ACE, Alu FX!llB, Alu PLAT, Alu Dl, and Alu PV92.
The number of chromosomes examined and allele frequencies for each five Alu insertion
polymorphic loci are given in Table 4.23. Locus specific allele frequencies for the seventy
one world populations studied are presented in Table 4.24. All the loci were polymorphic in all
populations, with the lowest level of polymorphism at Alu FXIIIB (0.0 15) in African Pygmies
followed by Toto (0.033) in Alu PLAT.
Heterozygosity for each locus for each population are given Table 4.25. All populations
exhibited high levels of diversity at most of the loci. The heterozygosity for Alu FXI!IB was
relatively low in all populations. The average heterozygosity and associated stadard error for
each locus for each 71 populations of world are given in Table 4.26. The average heterozygosity
over all five insertion polymorphic loci for each population was substantial and varied from
23% (Australian) to 50% (Iyenger).
The GsT value for each locus (Table 4.27) was calculated in order to measure the degree
of gene differentiation between these populations. They ranged from a low of 0.078 for Alu
D I to a high of 0.215 for Alu FXlllB. The GsT value for all loci was 0.128. which revealed
that 0.2% of the total variation in allele frequency differences at these loci \\as due to
differences between populations and 99.8% was due to differences within populations.
l2H
fllhk .S.2H ( ;t•nt·lk Uhlam·~ uf sn~nl' unc Wurld pnpulatiuns
', r~u 1 / b 8 9 10 11 11 13 14 15
\J UUI'!l 1J U.\1(1 0 O_H)f () 0444 0 04 t 0 01J4 () 0/~8 0 01/4 0 Oltil 0 0062 0 0/~~J 0 0;158 0 0096 0 0059 0 0033 0 04~0 0 0503 0 0093 0 0031 0 0116 0 0063 u 0403 0 0311 0 0115 0 0014 0 0091 0 01 0 0094 ,, 0 0864 0 0939 0 037 0 0174 0 0334 0 0315 0 0139 0 0247
10 UOB99 0091 00409 0 0181 00333 0 0316 0 0153 0 0259 0 0003 11 0 1188 0 1341 0081 0 0644 00885 00768 00554 0 0685 00263 00283 1) 0 1409 0 1531 0 0964 0 0671 0 0864 00792 00574 0.0776 0 0227 0022 0008 1:1 0 14/9 0 1502 01004 0 0689 0 0926 0 0808 0 0586 0 0777 0 0295 0 0293 0 0064 0.0045 14 01173 0 123 0 0557 0 0341 0.0576 0 0504 0 0256 0 0435 0 0048 0 0054 0.0165 0 0133 0.0157
" 0 1934 0 2112 0 1444 0 1086 0 01263 0 12 0.0948 01261 0 0504 0.0474 0.033 0 0101 0 0175 0 0348
16 0 1146 0 129 0 0797 0 05 0 062 0 0601 0.0444 0 06 0 0149 0.0134 0.0155 0 004 0 0129 00137 0 0167
17 01304 0 1374 0 0736 0.0463 0 0705 0 0643 0 0387 0.0546 0.0091 0.0093 0 0095 0 0061 0 0092 0.0023 0 0252
18 0 1159 0 12 0051 0 0373 0 0625 0 0499 0 0249 0.0472 0.0105 0.0124 0.0176 0.0188 0 0171 0 0027 0.0422
19 0 177 0 2029 0.1182 0 1077 0 1326 0.1199 0 0883 0 1213 00446 0047 00211 0.0163 0 0265 0 0288 0 0245
20 0 1111 0 1217 0.0647 0 0426 0 0617 0.0505 0 0345 0 0509 0 0086 00092 0007 0 0043 0 0091 0.0043 00246
21 0 0484 0 05 0.0094 0 0045 0 0092 0.0049 0 0013 0 0128 0.0201 0 021 0 0698 0.0678 0 0701 0.0342 0 1029
22 0 0509 0 0601 0.0136 0 0104 00105 0.0087 0 0061 0.0219 0.0226 0.0227 0.079 0 0696 0.0777 0 0388 00988 23 0 0447 0 0554 0.0109 0 0074 0.0084 0.0091 0 0057 0.0154 0.0173 0 0179 0.0726 0 0667 0.0774 0 0356 0.1088 24 0 0538 0 057 0.0191 0 0097 0 0088 0.0076 0 0077 00207 0 0276 0 0268 0.0855 0 0751 0.0799 0 0442 0.1037 25 0.0438 0 051 0.0112 0 0037 0 0069 0.0075 0 0036 0 01 0.0137 0.0143 0.0644 0.0611 0 0697 0.0311 00972 26 0.0435 0.0548 0 0109 0.0094 00108 00074 0.0042 0.0191 0 0162 0 0172 00612 0 0563 0 0642 0.0305 0 0878 27 0 0942 0.1091 0.0487 00284 0.0391 00378 00219 0.0406 0.0045 00035 00303 0 0169 0.0276 0.0081 0 0334 28 0 1207 0.1307 0.0573 0 0361 0.0577 0 0528 0 0274 0.0476 0.0045 00045 00233 0015 0 0214 0.0013 00328 29 0.1075 01192 0.0539 0 0318 0.0463 0.0428 0 0236 0.0452 0.0055 0 0044 0 0291 0 0152 0 0229 0.0053 00289 30 0 0585 0.0742 0.0326 0.02 0.0262 0 0275 0.0189 0.0246 0 0073 0.0082 0.0331 0.0313 0.0444 0 0196 0.0638 31 0.0547 0 0696 0.0293 0 0185 0.0248 0.025 0 017 0.0228 0.0075 0 0088 0 0321 0032 0 044 0 0196 0.0658 32 0.0871 0 1162 0.0399 0 0422 0.0504 0.0489 0 0322 0.0545 0.0178 0 0198 0.0541 0.0454 0 0655 0 027 0.0703 33 0.0618 0 0684 0 0333 0.0132 0.0206 0.0285 0.0204 0.0135 0.0135 00137 0.0593 0.0587 0.0715 0.0327 0.0988 34 0.086 0 1017 0.0673 0 0361 0.0392 0.049 0.0415 00407 0.1076 0.0155 0.0461 0 0341 0.0516 0.0326 0 0581 35 0.0994 0 1189 0.0839 00523 0.0565 0.0651 0.0562 00567 0.244 0.0225 0.0383 00275 0 0461 0.0361 0.0492 36 0.0477 0.053 0 0364 0.0513 0.0593 0.0412 00416 00503 0.0569 00632 00506 0.0764 0.0715 0.0627 01272 37 0076 0 0847 0.0727 0.045 0.0514 0.055 0 0509 0.0417 0.0289 00293 0.027 0.0325 0.0417 0.04 0 0681 38 0.0791 0 0988 0.0472 0.0304 0.0378 0.0434 0.0307 0.0359 O.D1 0 0102 0042 0.0353 0.0538 0.0234 0.0645 39 0.0378 0 0357 0.0232 0.005 0 0092 0.012 0 0122 0.0028 0 019 0.0197 0.0552 0 0615 0 0652 0.0376 0 106 40 00796 0 1068 0.0373 0.0369 0 044 0.0426 0 0279 0.0484 0 0141 0.0158 0.0473 0 039 0 0576 0 0234 0 064 41 00754 00983 0.0306 0.0262 0 0298 0.0318 0.0201 00394 0.0151 0.0151 0 0674 0.0527 0.0711 0.029 0.0758 42 0.1023 0.1295 0071 0.0544 0.0629 0 0675 0.0516 0.0615 00193 0.0197 0.0372 0 0288 0 0506 0.0284 0.0524 43 0.0449 0.054 0.0228 0.0079 0.0086 00102 00085 0.0143 0011 0.0107 0.0503 0 044 0.0532 0.0267 0 0748 44 0.0777 0.1062 0.0379 0 0401 0.0455 0.0415 00285 00534 0.0172 0.019 0.0462 0.0368 0.0541 0.0247 0.0588 45 0 0592 0 0796 0.0286 0.0281 0.0355 0 0309 0.0207 0.0349 0 0118 0.0144 0 031 0.0323 0.0447 0.0204 0.0648 46 0.0374 0 0348 0.0148 0.0077 0.0165 0.0159 0 0129 0.0036 0.0243 0.0274 0 0616 0.0762 0.0787 00424 01292 47 0.077 0.0884 0 046 0 0326 0.0461 0.0508 0.0369 0.03 0.0194 0 022 0.0475 0.0561 0 0704 0 0343 0 1015 48 0 0623 0.085 0.0325 0.0284 0.031 0 0277 0.02 0.04 0 013 0.0138 0.0394 0.0307 0.0444 0.0219 0.0537 49 00565 0.0652 0.0309 0.013 0.02 0 0252 00177 0.014 0 0096 0.0102 0 0464 0.0467 0.0588 0 0264 0 0853 50 00372 0 0411 0.0242 0.0123 0.0176 0.0155 0.0142 0.0113 0.0158 0.0178 0 0346 0.0436 0.0475 0 0293 0.0862 51 0.0649 0.0762 0.0516 0.0236 0024 0.0302 0.0278 0.0284 0 015 0.0131 0.0424 0 0326 0.045 0.0298 00582 52 0 0664 0 076 0.06 0.0634 00424 0 0444 0.0404 0.0347 00224 0.023 0 0245 0.0294 0.0383 0 0334 0.0649 53 0 0508 0 0653 0.0233 0.0131 0.0164 0 0167 00108 0.0203 0.0075 0.008 0.0421 0.0369 0.0486 0.0209 0 0671 54 0.0455 0.0578 0 0175 0 0101 0.0148 0.0141 0 0081 0 0155 0.0078 0 0092 0.0421 00413 0.0514 00216 0.0762 55 0.0528 0.0663 0.0253 0 014 0.0198 0.0211 0.0134 0 0185 0.0067 0.0077 O.D385 0.0371 0 0492 00204 0 0714 56 0 0482 0.0557 0 0227 0.0061 00086 0.0128 0.0092 0.0107 0.0105 0 0102 0.0559 0 0508 0.0612 0.0283 00849 57 0.053 0 0599 0.0289 0.013 00212 0.0251 0.0178 00122 0 0112 0.0126 0 0442 0 0489 0.0591 0.0276 0 0911 58 0.1075 0.1286 0 0544 0 0456 00609 0 0538 0 0324 0 0591 0.0107 0.0118 0.0241 0.0153 0.0264 0 0084 0 0315 59 0.1281 0.1449 0.0729 0 0661 0.0844 0.0684 0 0461 00824 0.0282 0.0298 0.0241 0 0168 0.0192 0.0162 0 026 60 0.0485 0.0573 0.0056 0.009 0 00134 0 0098 0.0034 0.0179 0.0194 0 0213 0.0703 0.0691 0.0759 0.0338 0 1054 61 0 0932 0 1103 0 05 0.0442 0.0565 0.0444 0.0291 00576 0.0169 0.0184 0.0206 0.0154 0 0202 0 0129 00323 62 0.0932 0.1056 0.0459 0.0253 0.0375 0 0355 0.0191 0.0368 0.003 0 0022 0 0285 0 017 0.026 0 0062 0.0352 63 0.159 0.1827 0 1063 0.0813 0 0985 0 0946 0.0686 0097 0.0273 0.026 0.0257 00073 0 0207 0 0187 0.0084 64 0.115 0 1338 0 0581 00485 0.0675 0.059 0.035 0.061 0 011 0 0127 00192 0.0135 0 0227 0 0064 00312 65 0 1169 0.1416 0 0744 00603 00702 0 064 0 0464 0 0764 0 0209 0.0205 00269 0.011 0.0233 0.0173 00176 66 0 1309 0 1437 0 0618 0 0486 00709 0 0627 0 0341 0 0596 0 0097 O.D\08 0 0235 00167 0 0231 0 0029 00334 67 0 0492 0.0459 0.0177 0.0239 0.0394 0 0273 0 0202 0 0205 0.035 0.0407 0 0513 0 0767 0 0718 0 044 0 1331 68 0.0945 00773 0 0428 00435 0.0721 00583 0 0426 0 0357 0 0591 0 0657 00774 0 1126 0 0991 0 0645 0 1783 69 0 1495 0.1426 0.0596 0.0716 0.1107 00902 0 0579 0 0736 0 0634 0 0711 00809 0 1095 0 0998 0 0576 0 163 70 0.1927 0 2173 01268 01002 0.1243 01214 0087 0 1156 0 0358 0 035 0 0339 0 0161 0 0316 0 0243 0 0167 71 0 1452 0 179 0 1102 0 0903 0.0943 0093 0 0771 0 1098 0 0421 0 0398 0 0494 0 022 0 0416 0 0399 0 0173
129
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1& 11 00083 18 0 0221 0.0073 19 0 0215 0 0218 0 0297 20 0 0048 0 0018 0 0085 0 0193 21 0.052 0 0493 0 0336 01018 0 0443 22 0.0508 0 054 0 0403 0 1009 0 0476 0 0026 23 0 0469 0 049 0 0395 0 0951 0 0428 0 0044 0 0024 24 0 0554 00597 00462 0.1162 0.0541 0 0031 00019 0 0064 25 00426 0 0432 00352 0.0922 0.0377 0 0037 0 0041 0.0007 0.0071 26 0 0401 0 043 0 0313 0.0823 0.0354 00034 0002 0.0021 0.006 0.0028 27 0 0082 0.0111 0 0151 O.D38 0.0086 0.0258 0 0228 0.0206 0.028 0.0191 0.0172 28 00132 0 0041 0.0058 0.0285 0 0061 0 0346 0.0359 0.0327 0.0423 00296 0 029 0.0051 29 0 0096 0.0087 0 0107 0.0359 0.0082 0 0278 0 0261 0.0259 0.0304 0.024 0.0213 0.0012 0.0029 30 0.0178 0 0222 0.0274 00512 0.0157 0.0249 0 0242 0 015 0.0319 0.0124 00147 0.01 0.0184 0 0166 31 0 0191 0.0226 0 0261 0.0516 0.0158 0.023 00232 0.0142 0.0308 0 0116 0 0133 0 0109 0.019 0.0173 0 0002 32 00338 0.0338 0.032 0.0431 0.0278 0.0361 0.0288 0.0215 0 0431 0.0239 0.0208 0.0154 0.0208 0 0209 0.0129 33 00393 0.0385 0.0446 0.0903 0 0361 0.0254 0.0272 0 015 0.0317 0.0111 0.0223 0.0237 0.0311 0 0309 0 0091 34 0 0162 0 0298 0.0494 0.0675 0.0252 0.0468 00428 0.0321 0.0472 0.0285 00352 0 0156 0.0288 0.0235 0.0098 35 0 013 0.029 0.0533 0.0555 0.0231 0.0637 0.0596 0.047> 0.0659 0.043 0.0482 0.0205 0.033 00289 0.0138 36 0.0738 0.0693 0.0514 0.088 0.0554 0.051 0 0629 0.0579 0 0708 0053 00455 00686 0 0727 00733 00466 37 0 0219 0.0325 0.052 0.0644 0.0249 0.0619 0.0667 0.0526 0.0713 0.0448 0.0511 0.0343 0.0437 0.0427 0 0161 38 0.0202 0.0246 0.0355 0.0522 0.0202 0.0368 0.0333 0 0215 0.0426 0.0196 0.0245 0.0116 0.0193 0.0189 0.0031 39 0.0436 0.0444 0.0441 01033 0.0391 0.0166 0.0234 0.0151 0.0235 0.0096 0 0178 0 0309 0 0407 0.0372 0 0142 40 00272 0.0293 0.0287 0.0412 00288 0.0319 0.0255 O.Q186 0.0383 0.0202 0.0172 0.0115 0 0182 0 0171 0 0087 41 0.0354 0.0391 00359 00672 0.0341 0.0198 0.0118 0.0083 0.0207 0.011 0.01 0.0116 0.022 0.0166 0.0133 42 0.0174 0.0249 0.0415 0.036 0.0194 0.059 0.0541 0.0409 0.0666 0.0392 0.0415 0 0167 0.0237 0 0245 0 0096 43 00262 0.0339 0.0333 0.0789 0.0271 0 0096 O.OOBB 00055 0.0108 0.0037 0.0051 0.012 0 0256 0.0175 0.007 44 0.0264 0.0308 0.0278 0.0386 00226 0.0318 0.0246 0.0204 0.0372 0.0226 0.016 0.0117 0.0199 0.0167 0 0117 45 0 0229 0.0245 0.0228 0.0399 00161 0 027 0.0261 0.0185 0 0371 0.0174 0 0145 0.0134 0.0198 0.0192 0.0041 46 0 0608 0.0525 0.0449 0 1075 0.0476 0.0189 0.0285 0 0181 0.0318 0.0128 0.0215 0.0425 0.0468 0.0485 0.0199 47 0.0429 0.0362 0.0447 0.0697 0.0334 0.0473 0.0514 00334 0.0616 0.0285 0.0393 0.0335 0.0343 0.0416 0.0114 48 0 0188 0.0271 0.0255 0 0447 0.0181 0.0226 0.0174 0.0146 0026 0.0153 0.0095 0 0076 0.019 0.0216 0.007 49 0.0298 00307 0.0366 0.0753 0.0266 0 0234 0 0253 0.0139 0.0305 0.01 0.0182 0.0179 0 0256 0.025 0.0039 50 0.031 0 0332 00328 0.0745 0.0251 00209 0 0273 0 0191 0.0308 0.0137 0.0169 0.025 0 0336 0.0312 0.0081 51 O.Q105 0.0295 0.0426 0.0716 0.0202 0.0314 0 0294 00227 0.0311 0 019 0.0228 0 013 0.0282 0.0199 0 0079 52 0 0186 0.0281 0 0434 0 0593 0.0202 0.0503 0.0545 0.0421 00594 0.0352 0.0399 0.0272 0.0366 0.035 0.0106 53 0.0215 0.0271 0.0269 0.061 0.0201 0 0136 0.0117 0.0066 0.0175 0 0054 0.0056 00081 0.0191 0.0138 0.0025 54 0 0266 0.0288 0 0261 0.0644 0.022 0.0117 0.0119 0.0058 0 018 0.0042 0.0053 0.0118 0.021 0 0175 0 0028 55 00224 0.0252 0.0274 0.0594 0.0191 0 0183 0 0182 00098 0.0249 0.0075 0 0104 0.0107 0 0193 0 017 0.0007 56 00315 0.0361 0 0367 0.0861 0.031 0.0109 O.D108 0005 0 0131 0.0028 0.0079 0 0148 0 0267 O.D208 0 0071 57 0.0332 0.0319 0.0364 0.0762 0 0275 0.0246 0.0285 0.0162 0.0341 0.0116 0 0201 0.0221 0.0283 0 0291 0 0049 58 0 0131 O.D108 0.0108 0.0177 0.0073 0.039 0.0361 0.0331 0.0467 0.0324 0 0262 0.0061 0.0059 0 0065 0 0156 59 0.0231 0.0185 0.0112 0.0174 0.0146 0053 0 0531 0.0566 0.0619 0 0554 0.042 0.0206 0 0169 0 0164 0.0399 60 0.0537 0.0494 0.0328 0.0914 00436 0.0028 0.0034 0.0027 0.0083 0.0035 00025 0 025 0 0327 0.0282 0.0209 61 0.0151 0 0158 0 0105 0.0236 0.0093 0.0348 0.0345 0.0356 0.0424 0.0343 00238 00112 0 0136 0.0107 0.0213 62 0.0089 0.0096 0.0125 0.0389 0.0078 0.0233 0 0218 0 0197 0.0265 00177 0.0163 0.0002 0.004 0.0009 0 0099 63 0 0088 0.013 0.0268 0.0122 0 0117 0 0768 0.0713 0.0682 0.0809 0.0662 0.0599 00154 0.0148 0 0144 0 035 64 0.014 0.008 0 0079 0.014 0.0057 0.0432 0.0425 0.0386 0.0533 0037 0.0313 00087 00051 00082 0 0181 65 0 0085 0.0162 0.0214 O.D185 00106 0 0516 00454 0 0452 0 0545 0 0449 0 0353 0 0086 0 014 00093 00234 66 0.0185 0 0062 0.0047 0.0204 0.008 0.0422 0.0435 00406 0 0524 0 0382 0 035 0 01 0 0018 0 0068 00246 67 00771 00546 0.0369 0 094 0.0483 0.0294 0 0436 00357 0 0498 0 0299 0 0318 0 0554 0 0527 0 0585 00339 68 0.1115 00776 00556 01327 0.0786 0.0548 0.0772 00676 0 0812 0 0588 0.067 0.0915 0 0768 0 0907 0 0701 69 0.1173 0.0729 0.0444 01019 00777 0 0703 0.0885 0 0822 0 099 0 077 0.0788 0 0909 0 0659 0 0855 0 0851 70 0 0205 0 0174 0 0339 0.0115 0.0199 00977 0.0927 00863 0 04049 0 0839 0 0803 0 0266 0 0193 0 0245 0 0472 71 0.0163 0.0351 0 0495 0 0306 0.0282 0 0804 00674 00673 0 0771 00689 0 058 0 021 0 0327 0 0235 0 0403
130
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10 11 17 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 0 0134 33 0.0099 0 0288 34 0 0122 0 0313 0.0137 35 0.0164 0 0358 0 0246 0.0028 36 0.0415 0 0675 0 0702 0.0867 0 0887 37 0.0165 0.0521 0 0264 0.0148 O.D108 0.0576 38 0 0046 0 0115 0.0089 0.0065 0.0102 0.0699 0.021 39 0.0131 0.0453 0 0081 0.0248 0.0366 0.0468 0 0246 0.0235 40 0.0091 0 0006 0 025 0.0254 0.0294 0.0609 0 0436 0.0087 0.0385 41 0.0141 0.0063 0.0208 0 0258 0.0365 0 0768 0.0559 0.0128 0.0336 0 0056 42 0.0117 0.0139 0.0247 0 0111 00083 0.0795 0 0223 0.0044 00435 0.0114 0.0231 43 0 0068 0 0243 0 0114 0.0159 0.0263 0.0513 0.029 0 0141 0.0084 0.0187 0.0127 0.0289 44 0.0116 0.0024 0.0333 O.D305 0 0388 0.0562 0.0476 0 0143 0.0431 0.0013 00074 0.0159 0.0197 45 0 0032 0.008 0.0219 0.0243 0.0265 0 0337 0.027 0 01 0.0252 0.0051 0.0145 0.0137 0 0138 0 0054 46 0 0178 0 0454 0 0123 0.0417 0.0545 0 0383 0 0365 O.D305 00048 0.0405 0.0387 00521 0.0182 0.0463 0.0261 47 0.0118 0.0267 0 0089 0.0226 0 0267 0.0626 0.0226 0.0099 0.0213 0 0244 0 0332 0.0176 0.0282 0 0341 0 0186 48 0.0068 0.0085 0.0269 0.0222 0 0263 0.0472 0.0352 00127 0.0296 0 0050 0.0084 0.0172 0.0096 0 0029 00043 49 0.0043 0.0232 0.0013 0.0106 0.0188 0 0571 0.0187 0.0057 0 0068 0.0189 0.0184 0 0184 0.0079 0.0253 0.0138 50 00065 0.0654 00142 00237 00295 0.027 0.0151 0/0194 0.0048 0 0286 0 0326 0.0321 0.0091 0 0304 0.0126 51 0.0091 0.033 0.0139 0.0035 00083 0.0686 0.0141 0.0107 0.0153 0.0258 0.0239 0.0189 0.0069 0.0281 0.0203 52 0 0106 0.0427 0.0218 0.013 O.D106 00483 00008 0.0165 0.0192 0.0347 0 0457 0.0192 0.0214 0 0377 0.0192 53 0 0023 0 013 0.011 0 0143 0.022 0.0469 0.0265 00077 0.0128 0.0088 0.0078 0 0182 0.0021 0 0100 0 0058 54 0 0021 0.0144 0 0096 0 0184 00267 0.0399 0.0268 0 0093 0.0098 00104 0.0102 0.0214 0.0030 00122 0 00053 55 0.0006 0.0135 0.0067 0 0122 00188 0.0458 0.0204 0.0046 O.D105 0.0095 0 0111 0.0143 0.0045 0.0126 0 0051 56 0 0072 00249 0 0052 0 0145 0.0265 00586 0 0296 0.0117 0.0060 0 0200 0 0127 0.0283 0 0015 0.0237 0 0165 57 0.0048 0.0261 0.0023 0 0148 0 0225 0.0502 0.0175 0 0083 0.0057 0.0261 0.0231 0 0212 0.0102 0 0281 00137 58 0 0158 0 0105 0.0392 0 0315 0.0319 0 0603 0 0438 0.0180 0.0485 0 0085 0 0177 0 0169 0.0265 0 0072 0 0103 59 0.0387 0 0348 00746 0.0629 0.0614 0 0588 0.0674 0.0495 0.0732 0 0314 0 0433 0.0468 0.0472 0 0246 0 0281 60 0 0191 0 0234 0.0239 0047 00624 0.0480 0.0633 0 0306 0 0209 0.0212 0.0127 0.0506 0 0117 0 0217 00193 61 0 0201 0.0225 0.0509 0.0419 0.0421 00422 0.0456 0.0312 0.0480 0.0183 0 0277 0 0316 0.0263 0.0129 0 0131 62 00106 0 0167 0.0224 0.0169 0.0224 00656 0.0340 00123 0.0284 0.0127 0 0126 0 0187 0 0115 0 0132 0 0135 63 0.037 0.0332 0.0639 0 0356 0 0296 0.1019 0.0500 00323 00788 00298 00421 0 0225 00507 0.0282 0 0343 64 0 0181 0 0139 0.0418 0.0354 0.0349 0.588 0 0439 0 0208 0 0508 0 0120 00235 0.0190 0.0313 0.0112 00122 65 0 0242 0.0211 0.0541 0 0322 00295 0.0731 0.0456 0 0264 0.0610 0.0172 0.0265 0.0213 00321 0 0129 0 0192 66 0 0248 0.0204 0.0422 0 0408 00433 0 0718 0 0538 0 0260 00528 0 0190 0.0265 0 0277 0 0354 0 0197 0 0212 67 0 0297 0 0541 0.0386 0.0711 0.0803 0.0142 0 0523 0 0513 00237 0 0497 0 0571 0 0692 0 0366 0 0515 00287 68 00654 00941 0.0621 0 1115 01246 0 0461 0 0884 0 0873 0 0469 0 0906 0 0958 0 1118 0 0729 0 0964 0 0675 69 00808 0081 00878 01356 0.1466 0.0678 0 1252 0 0971 0 0861 0 0827 00925 01140 0 0962 0 0873 0 0717 70 0 05 0 0396 00736 00473 0 0401 0 1245 0 0641 0 0395 0 0973 0 0382 0 0537 0 0270 0 0704 0 0396 0 0466 71 0 0427 0 0349 0 0749 0 0376 0 0321 0 1124 0 0602 0 0384 0 0883 0 0309 0 0393 0 0276 0 0500 0 0258 0 0389
131
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" ;I 10 11 1? 13 14 1; 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 0.0185 48 0.0371 0.0333 49 0 0115 0.0074 0.0187 50 0.0084 0.0187 0.0191 00084 51 00323 00275 0.0161 0.0095 00140 52 0.0298 0.0195 0.0263 0.0140 0.0097 0 0106 53 0.0196 0 0208 0.0041 0 0064 0.0093 0.0084 0.0188 54 0 00133 0.0175 0 0065 0.0053 0 0065 00118 0 0189 0.0008 55 0 0151 0 0119 0 0076 0.0027 00 70 0.0088 0.0140 0 0013 0.0010 56 0.0138 0.0213 0.0146 0.0040 0 01a1 0 a083 0 a226 O.Oa35 0.0035 0.0040 57 0.00080 0.0057 a.0214 a.0006 0.0062 0.0128 00130 00083 0.0060 0.0036 0.0062 58 0.0536 0 0373 0 0090 0.0301 0.0338 0.0301 0.0356 00164 0 0190 0 0180 0.0309 0.0326 59 0 0788 0.0734 0.0240 0.0610 0.0520 00545 0 0574 00377 0 0404 0.0426 0.0572 0.0626 0.0092 60 0.0188 0.0395 0 0175 0 0216 00222 0.0349 0.0511 0.0114 0.0090 0.0150 0.0121 0 0225 0 0323 0 0496 61 0.0544 0.0523 0.0098 0.0387 0.0301 0.0330 0.0363 0.0191 0 0212 0.0233 0 0350 0 0401 0 0052 0 0040 0 0320 62 00389 0.0322 0.0086 0.0169 0.0230 0 0135 0.0268 0.0080 0.0111 0.01a3 a.a139 a.0205 0.0066 0.0203 0.0232 63 0.0940 0.0613 0 0288 0.0532 0.0612 0.0396 0.0454 0.0397 o.a465 a.0413 a.0566 0.0581 0.0108 0 0176 0 0726 64 0.0542 0 0366 0 0134 0 0324 0.0350 o.a346 o.a361 a.0204 a.o222 0.0208 0.0353 0.0340 a oa07 0 0085 0 0364 65 0 0742 0.0563 0.0117 0 0425 0 0435 o.a295 0.0382 00233 00290 a 0276 0 0402 0 0468 0 0047 0 0095 0 0476 66 0.0556 0 a4a4 a.0219 0.0352 0.0416 00404 00457 00258 0.0273 0.0260 0.0374 0 0372 0 0045 0 0119 0 0374 67 0.0113 0 0350 oa447 00328 0.0167 00574 0 0434 0 0340 0.0253 0 0299 0 0363 0 0263 0 0539 0 0646 0 0269 68 00270 00567 o.a902 00604 0.0466 0 0984 0 0800 0 0714 0.590 0 0636 00674 0 0514 0 0911 0 1026 00544 69 0.0600 00736 00934 0.0844 0 0786 0 1278 0 1133 0 0860 0.0748 0 0799 0 0918 0 0772 0 0757 00810 00616 70 0.1089 00639 0 0452 00643 0 0796 0 0574 0 0604 0 0557 0.0621 0 0544 007H 00691 00179 0 0287 0 0901 71 0 1096 00801 0 0250 0.0628 0 0699 0.0390 0 0543 0 0406 0 0504 00467 00595 0 0702 0 0196 0 0280 0 0758
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133
I hl' all~.:lc: frclJUCI\(~ l'(lr ca...:h ln..:u' in ca\:h population wa.., u~cd to calculate the Nci 's
... r.1ndard dl ... tann:' (Nci. I"J72) hctv.ccn the population~ lahlc 4.2H. Ba"cd on this di~tancc
· · 1 " 1 k, 1 l·t· 1 · indudin" the 26 ethnic mat! 1\, a 1\Ct!-!,hhor-JtHI\\Il~ ( NJ) trcc rclatmg llC ).1 g t)p,\ pnpu cl 101 s. t:"
pnpulatlllll' of India 1'0 prc..,cntcd in l"igurc ·l.\3.
PNGhHi<thlandot>!S Aus'tr-alta.ns
Figure 4.13 Neighbour-joining tree depicting genomic affinities among 71 global populations based on allele frequencies at 5 Alu insertion loci
On the tree, all the African populations clustered together, as did the East Asia and
Europe populations.ln the present study out of the three populations of Bastar, Bison-horn
Maria (BHR) and Muria (MUR) clusterd together, showing significantly nearer except
Abujhmaria (ABM). Bison-horn Maria as well as Muria lie in between the East Asian
populations, indicating that the Bison-horn Maria and Muria have similar genetic composition
with the East Asian populations.
It is seen that, by and large, the Indian populations lie in between the Mongoloid population
groups (Filipino, Malaysian, Koreans etc) and Caucasoid populations groups (Greeks. French.
134
"'~ ,.., ... ) II"".' Hknt tromthc dcn(.h1~ram that BP·am-horn Mana and Muria ... howing intcrmcdiatc
.tlltlltl_l \\tilt tllollgnlnul pnpulattnll'- .African populations an: genetically quite distance from
the <tlltcr global populations.Sincc the ancestral stale (non-insertion) nf these Alu
pnl_1 morphisms arc known, it is possible to root the unrooted NJ tree hy using the ancestral
population (with 1crn inscrtinn-alklc frequencies at allloei) as an outgroup. The root lies
l'ery dose tot he duster of African populations. This provides additional evidence for the
suppnrl oft he ··out-of-Africa" hypothesis of modern human ( Cann ct al1987; Bowcock ct
al. IIJIJ4).
Table 4.30 l>istance from Centroid & Heterozygosity among the population
Tribe l>istance from Centroid Heterozygosity
Ahujhmrnia 0.0100 0.4160 --
Bison-horn Maria 0.0142 0.3786 f---
Muria 0.0052 0.3949
Halba 0.0100 0.4269
Kamar 0.0052 0.4134
Cllinda Bhunjia 0.0031 0.4032
ChaukatiaBhunjia 0.0229 0.4069
Munda 0.0037 0.3991
Lodha 0.0577 0.3659
San tal 0.0059 0.4001
Katharia Tharu 0.0130 0.3494
RanaTharu 0.0130 0.3520
Dangwia(B) 0.0059 0.3847
Dangwia 0.0264 0.3312
Pachchimaha 0.0212 0.3752
Toto 0.0241 0.3360
Mizo 0.1319 0.2998
Ho 0.0241 0.3523
Tipperah 0.0123 0.3550
Badaga 0.0142 0.3977
lrula 0.0031 0.4202
Kota 0.0123 0.3608 l -
-- - -- --- ------- --~---
\unha ().(} 156 0.1H07 -
lod a 0.1 h~4 0.1141 -- ------ --- c--· - .
1da Rcddi 0.0017 0.407 5 - ------- ·-- --- -·-- -- --------- - --- ---- ----
·a Dora O.OOH3 0.1800 - ------ - ---- - -------·--
lll 0.0242 0.3770
Min 0.0031 0.2998 ---- ---------
M:Lx 0.1624 0.4269 -
4.10 Gene Flow Among Populations
With the objectives to know what are the genetic characteristics of the populations
under study and what are the evolutionary forces generated the observed patterns; and to
test whether in a group of incompletely isolated populations distributed over geographical
space (Wright's island model), the observed patterns of genomic diversity are generated by
interactions with populations outside the set of populations under consideration, Harpending
and Ward (1982) derived a regression of heterozygosity on genetic distance. This theory
assumes that the genetic distance of an island population from the gene frequency centroid
(the overall mean gene frequencies of all populations) and the relative homozygosity of that
island population should be linearly related if exchange with populations from outside is the
same for each island. If gene flow from outside varies in amount from one population to
another, this linear relationship no longer holds. Very isolated populations should be less
heterozygous than the linear prediction, whilst populations which receive more genes should
be more heterozygous than predicted. As this model states that given the uniform systematic
pressure (gene flow) from outside, the average genetic heterozygosity within a population is
negatively correlated with genetic distance from the centroid of the gene frequency array
(rii), higher than average heterozygosity than predicted by the regression model and lie above
theoretical regression line. Conversely, population experiencing isolation and less than average
gene flow from outside show lower values of heterozygosities and lie below the regression
line.Harpending and Ward (1982) model is somewhat analogous to the Wright's island model.
We have performed this centroid analysis hy pooling the data of the present study with
those of other populations oflndia. This analysis was hased on data of eight Alu polymorphic
loci.The result of heterozygosities along with descriptive statistics. at eight individual loci i.e.
Alu mt NUC, Alu APO, Alu ACE,Alu CD4, Alu PV92, Alu Dl, Alu PLAT, Alu FXlllB are presented in Table 4.15
A regression plot of average heterozygosity verses distance of each of the populations
under study along with 23 populations taken for comparison, to the centorid is given in
Figure 4.14, along with theoretical regression line. From the Figure 4.14 it is indicated the
136
4400 H<1lba.
.4200; lrula
~IIJhmana ar KondaRectflaukatraBhu
.4000 1 Ch'~f£'' dag~
M...wa ~
>- I Dangl!na-B f-.3aooJ
KoY@fl<i«l,na Kurumba (/) -' PanBhlrnaha
0 I Lodha CJ ' >- Kota
N .3600 j T1~nli 0 lf.ft\~c .rw,.,o 0:: IJ..I 34ooJ Toto f- Danguna IJ..I I
.3200- Toda
Mrzo
.30001
.2800 , --- --,-··------,--- -~ --- - 1 --··- ~----------.---- - -- --r--~r - ---------j
- 0250 0.0000 .0250 .0500 .0750 .1000 .1250 .1500 .1750
DISTANCE FROM CENTROID
Figure4.14: Plot of heterozygosity vs distance from allele frequency centroid of twenty six Indian ethnic populations based on allele frequency data of eight loci
137
11.111 lhl· Jhi(HJIHilllll' ha\~ c.\(H.:ncnccd lc,~cr ~cnc flu\\ than prcdidcd. \\hilc the other
pt~pul.tttorh ha\c ht~hcr than prcd11..:tcd I he fi~urc abo indi~:tHcs that for most of the populations
'"''"""~ g""') i' hi!:htr hut hu<l 11 n11rrow range (O.JJOO lo 0.4.100) 11nd lend lo dnslcr close
"''" apr"'imately 1UlU25 ro 0.00225 di,tance from centroid. Only three populations vii. l.odha.
\111o and Jod11 seems t<><tuit distinct and fall awny from the ccntroid.The analysis revealed
that the i\BM & MliR have experienced higher gene !low than predicted in terms of having
""'n than predicted heterozygosity while the IHIR has experienced less gene !low than
l"'·diclcd. In the regression plot i\hujhmaria is far moved above the centroid line. which
ma) suggest that they have experienced more external gene flow than BH Rand MUR. BHR
is plnced nearer to theoretical regression line as well as relatively closer to the centroid. This
observation may be explained by population history. A likely explanation is that a gene l1ow
occurcd prior to the sub-division of these populations into largely endogamous units, long
before the region has been restricted. However, evidence favouring inflow of genes from
difi'crent external sources pertaining to the study populations, while scanning through
anthropological literature. Although Jain (1961) reported that extensive tour of that area
was made by the foreigners since 1863 and that might be the source ofthe inflow of genes.
The ethnohistory of ABM suggest that they have migrated from Karnataka, South India to
Bastar around 1112 AD (Dubey,2000) and after they were driven away with their kinsmen.
Therefore, their position in the above regression plot may reflect bottleneck effect of certain
degree.Abujhmaria inhabit in geographically isolated area- Abujhmarh, and the region is a
restricted area for about 25 years by the Government order.Even one has to take permission
from the Government authority to enter Abujhmarh. However, some morphological evidences
suggest intermarriage with some Aryan stock (Guha, 1931 ).
Noronha (1962) also states that there are 3 district racial type in Abujhmarh-
(1) the red bronze, almost cauiasion type,
(2) very dark symmetrical pravidian type and
(3) the primitive Bushman type. He further noted that the orignal population in
Abujhmarh must have been the Bushman type who was squeezed into the centre of
Marh by sucessive waves of outsiders of the two other types. He estimated that
the Bushman type is 20% and the Caucasian type and Dravidan type are 40%.
each. However, it is difficult to understand how these have been coupled, as these
are quite distinct and extreme types. Noroha opines that this area was orurrun by
the Aryans in the 4th century A.D. as is recorded in the Puranas.
The history of Abujhmarh in particular and that ofBastar in general. still remains lar\:t:h
unexplored. Possibly one reason for this has been the widely held belief that the AB'Vl arc
the most primitive tribe, as popularised in anthropological I iteraturc. acwrdi n\:h to Duhc'. ' . .
13K
tilt· \11\h are lit> dt>uht a !!L't>}:!itphi.:all) i,o{att:d art: c<:onomicall) backward pcopk but
thn .ue nu m..:ans !'rum the trihal ,tock. I ht:) art: the de<:.:mknts of an aristnnati<: dill:
,·I·•" "hi .:It in th.: days of) llrl' ruled llastar region through a powerful ( 'hakrakote Kingdom
11 11h "'capital loeated at llarsur. I k was also quit<: ama1.<:d by th.: fair compkxion, blu.:
cH·.cvtn:mcly handsom, sharp katur.:s and dirty rust ~:nlnured hair of a good proportion of
tlt<·nt. Sueh feature do not suppnrt the views tlwt th<.: II ill Marias ar<: tribes like Oonds,
1\aigas. 1\.orkus and So on (Dubey. 2000).
tilasl'urd ( 1906) during his visit to 1\hujhmarh recorded that their phisique is good,
though most arc small and rarely is a dark coloured Maria seen. Their skin arc wonderfully
fair. in snmc cases as light in tint as that of a fair Brahman.
The ABM still retain thccomplexion and feature of their elite ancestors, who originally
came from Kannad country of the Tamil Nadu. These observation suppor our genetic findings.
Further, it was found that all the poulations showed considerably higher levels of
heterozygosity than that predicted by the Harpending- Ward gene flow model.lfthis pattern
of high heterozygosities were simply due to higher levels of gene flow, then one would have
expected that tribal populations ofBastar would be genetically less differentiated. The findings
reveals that the co-efficient of gene differentiation among tribes ofBastar is lower than that
among populations inhabiting all other parts oflndia. Drift effect could have accentuated the
process of genetic differentiation and has altered ancestral gene frequencies. Several lines
of evidence suggest that genetic drift/gene flow has been the major evolutionary force to
shape genetic variation in populations. This, by and large, is in agreement with anthropological
findings. Since this analysis does not permit timing of the period during which the gene flow
may have occured between these populations, we unable to offer a clear interpretation of
this finding. This represents important feature of populations under study, which has to be
taken into account in any attempt to reconstruct the history of this populations.
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