bogus claims about aryan invasion

8/12/2019 Bogus Claims About Aryan Invasion

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Brief Comm unication 1 3 3 1

D e e p c o m m o n a n c e s tr y o f In d i an a n d w e s t e r n E u r a s ia nm i t o c h o n d r ia l D N A l in e a g e sT . K i v is i ld * , M . J . B a m s h a d r, K . K a l d m a * , M . M e t s p a l u * , E . M e t s p a l u * ,M . R e i d l a * , S . L a o s * , J . P a r i k * , W . S . W a t k i n s t , M . E . D i x o n t , S . S . P a p i h a * ,S . S . M a s t a n a , M . R . M i d ] , V . F e r a k a n d R . V i l l e m s *A b o u t a fi ft h o f t h e h u m a n g e n e p o o l b e l o n g s l a r g e l ye i t h e r t o I n d o - E u r o p e a n o r D r a v id i c s p e a k i n g p e o p l ei n h a b i t in g t h e I n d i a n p e n i n s u l a. T h e C a u c a s o i d s h a r ei n t h e i r g e n e p o o l i s t h o u g h t t o b e r e l a t e dp r e d o m i n a n t l y t o t h e I n d o - E u r o p e a n s p e a k e r s .A c o m m o n l y h e l d h y p o t h e s i s , a l b e i t n o t t h e o n l y o n e ,s u g g e s t s a m a s s i v e I n d o - A r y a n i n v a s io n t o I n d i a s o m e4 ,0 0 0 y e a r s a go [ 1 ] . R e c e n t l im i t e d a na ly s is o fm a t e r n a l l y i n h e ri te d m i t o c h o n d r i al D N A ( m t D N A ) o fI n d i an p o p u l a t i o n s h a s b e e n i n t e rp r e t e d a s s u p p o r t i n gt h is c onc e pt [ 2 ,3 ] . H e r e , t h is in t e r p r e t a t ion i s que s t ione d .W e f o u n d a n e x t e n s iv e d e e p l a t e P l e i s t o c e n e g e n e t i cl i n k b e t w e e n c o n t e m p o r a r y E u r o p e a n s a n d I n d i a n s ,p r o v id e d b y th e m t D N A h a p l o g r o u p U , w h i c he n c o m p a s s e s r o u g h l y a . fi ft h o f m t D N A l i n e a g e s o f b o t hpopu la t ions . Ou r e s t im a t e f o r t h is s p l i t i s c los e t o t hes u g g e s t e d t i m e f o r t h e p e o p l i n g o f A s i a a n d t h e f i r s te x p a n s i o n o f a n a t o m i c a l l y m o d e r n h u m a n s i n E u r a s i a[ 4 - 8 ] a nd l i k e ly p r e - da t e s t he i r s pr e a d t o E ur ope . On ly as m a l l fr a c ti o n o f t h e C a u c a s o i d - s p e ci f ic m t D N Al i n e a g e s fo u n d i n In d i a n p o p u l a t i o n s c a n b e a s c r i b e d t oa r e la t i v e ly r e c e n t a dm ix t u r e .Addresses: *D epartment o f Evo lutionary Biology, Tartu Universi ty,Riia23, Tar tu 51010, Estonia. Depar tments of Pediatr ics and HumanGenetics, Eccles Insti tute of H uman Ge netics, Universi ty of U tah, SaltLake City , Utah 841 12, USA. *Depar tment of Human G enet ics ,Universi ty of New castle-upon-Tyne,New castle, UK. Dep artment ofHum an Sciences, Loughborou gh Un iversi ty, Loughb orough, UK.Veterinary Col lege o f Srinagar, Kashmir 190003 , India. YFacultyo fNatural Sciences, Cemen ius Universi ty, 842 15, B ratislava, Slovakia.Correspond ence: R. Vi llemsE-mail : rvi llems@e bc.eeReceived: 10 Aug ust 1999Revised: 28 S eptem ber 1999Accepted: 29 Septem ber 1999Published: 8 Nove mb er 1999Current Biology 1999, 9 : 1 3 3 1 - 1 3 3 4096 0-98 22/9 9/ - see f ront matter 1999 Elsevier Science L td. Al l r ights reserved.

e s u l t s a n d d i s c u s s io nT h e r e c e n t A f r ic a n o ri g in o f m o d e r n h u m a n s i s n o w s u p -p o r t e d b y p a l a e o a n t h r o p o l o g i c a l , a s w e l l a s s e x - s p e c i f i ca n d a u t o s o m a l g e n e t i c , e v i d e n c e ( f o r re c e n t r e v ie w s , s e e[ 8 , 9 ] ) . T h e c o n c o r d a n c e b e t w e e n t h e i n t e r p r e t a t i o n o fd a ta o b t a i n e d b y m t D N A , Y - c h r o m o s o m a l a n d m o s t o f t he

a u t o s o m a l m a r k e r s i s e n c o u r a g i n g a n d s u g g e s t s t h a t , i r r e -s p e c t i v e o f t h e d i f f e r e n c e s i n t h e m o d e o f in h e r i ta n c e ,t h e s e t h r e e g e n e t i c a p p r o a c h e s p r o d u c e c o n s i s t e n t o v e r a l lf i n d i n g s i n t h i s c e n t r a l i s s u e .W e s e q u e n c e d t h e m i t o c h o n d r i a l h y p e r v a r i a b le r eg i o n I( H V R I ) a n d p e r f o r m e d e x t e n s i v e r e s t r i c t i o n f r a g m e n tl e n g t h p o l y m o r p h i s m ( R F L P ) a n a ly s is o f 5 5 0 I n d i a nm t D N A s a m p l e s . W e i n f e rr e d a p a r s i m o n i o u s p h y l o g e n e t i ct r e e f r o m t h e d a t a u s in g t h e m e d i a n n e t w o r k a p p r o a c h[10] , which i s par t icu la r ly su i tab le for in t raspec ies ana lys iso f m t D N A l i n e ag e s a n d o t h e r h i g h l y v a r i ab l e d a ta s e t s .F i g u r e 1 i s a n o u t l i n e o f th i s I n d i a n m t D N A t r e e w i t h int h e b a c k g r o u n d o f t h e p r e v i o u s l y d e f i n e d g l o b a l m t D N Al i n e a g e c l u s t e r s ( h a p l o g r o u p s ) [ 1 1 - 1 3 ] . C o n s i s t e n t w i t h t h er e c e n t o u t - o f - A f r i c a m o d e l o f h u m a n o r i g in s [ 1 4 ], a ll o f t h eI n d i a n m t D N A l i n e a g e s w e i n f e r r e d c a n b e s e e n a s d e r i v -i n g f r o m t h e A f r i c a n m t D N A l i n e a g e c l u s t e r L 3 a ,d e s c r i b e d i n [1 5 ]. W e f o u n d t h a t m o r e t h a n 8 0 o f t h eI n d i a n m t D N A l i n e a g e s b e l o n g t o e i t h e r A s i a n - s p e c i f i ch a p l o g r o u p M ( 6 0 .4 ) o r w e s t e r n - E u r a s i a n - s p e c i f i c h a p -l o g r o u p s H , I , J , K , U a n d W ( 2 0 . 5 ) , w h i l e t h e r e m a i n i n g1 9 .1 o f l i n e a g e s d o n o t b e l o n g t o a n y o f t h e p r e v i o u s l ye s t a b l i s h e d m t D N A h a p l o g r o u p s ( T a b l e 1 ) . W e n o t e t h a th a p l o g r o u p K s h o u l d n o w b e c o n s i d e r e d a s u b - c l u s t e r o fhaplogroup U [13] .T h e f i rs t a n d t h e m o s t p r o f o u n d l a y e r o f o v e r l a p b e t w e e nt h e w e s t e r n - E u r a s i a n a n d t h e I n d i a n m t D N A l i n e a g e sr e l a te s t o h a p l o g r o u p U , a c o m p l e x m t D N A l i n e ag e c l u s t e rw i t h a n e s t i m a t e d a g e o f 5 1 ,0 0 0 - 6 7 , 0 0 0 y e a r s [ 16 ]. U n t i ln o w , t h i s h a p l o g r o u p h a s n o t b e e n r e p o r t e d t o o c c u r i n iI n d i a n o r e a s t o f I n d i a a n d w a s c o n s i d e r e d a w e s t e r n -E u r a s i a n - s p e c i f i c h a p l o g r o u p . S u r p r i s i n g l y , w e f o u n d t h a th a p l o g r o u p U i s t h e s e c o n d m o s t f r e q u e n t h a p l o g r o u p i nI n d i a a s i t i s i n E u r o p e ( T a b l e 1 ) . N e v e r t h e l e s s , t h e s p r e a do f h a p l o g r o u p U s u b c l u s te r s i n E u r o p e a n d I n d i a d i f f er sp r o f o u n d l y ( F i g u r e 2 ) . T h e d o m i n a n t s u b c l u s t e r i n I n d i a isU 2 . A l t h o u g h r a r e i n E u r o p e , t h e S o u t h - A s i a n f o r m d i f f e r sf r o m th e w e s t e r n - E u r a s i a n o n e : w e s t e r n - E u r a s i a n U 2i n c l u d e s a f u r t h e r c h a r a c t e r i s t i c t r a n s v e r s i o n a t n u c l e o t i d epos i t ion (np) 16 ,129 [12] , which i s absent in Indian U2v a r i e t i e s ( F i g u r e 2 ) . W e c a l c u l a t e d t h e c o a l e s c e n c e a g ee s s e n t i a l l y a s d e s c r i b e d i n [ 1 5 , 1 7 ] a n d e s t i m a t e t h e s p l i tb e t w e e n t h e I n d i a n a n d w e s t e r n - E u r a s i a n U 2 l i n e a g e s a s5 3 , 0 00 + 4 ,0 0 0 y e a r s b e f o r e p r e s e n t ( B P ) . W e n o t e t h a t U S ,t h e m o s t f r e q u e n t a n d a n c i e n t s u b c l u s t e r o f h a p l o g r o u p U


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1332 Current Biology Vol 9 No 22

Figure 1

Rc

Current iologyThe skeleton network of Indian lineage clusters on the back ground ofcontinent-specif ic mtDN A haplogroups. Red, Indians; green, westernEurasians; yellow, eastern Eurasians; blue, Africans. H aplo gro upfrequencies are proportional to node sizes. All Indian, eastern-Eurasianand western-Eurasian mtDN A lineages coale sce finally to the Africannode L3a. The former are shown magnif ied to accou nt for highermtD NA divers ity in sub-Saharan Africans. The most likely root of thetree [15] is indicated within a pan-African cluste r L1. The dash ed lineleading from the African external node L3a to the Eurasian mtDNAvarieties identifies the position of L3a in the magnified part of the tree.

in Europe, has an a lmost ident ical coalescence age es t i -mate [13]. S ti ll , despi te their equal ly dee p t im e depth , theIndian U2 has not penetra ted western Euras ia , and theEuropean U5 has a lmost not reached India (Table 2) .Subclus ter U7 (among U* in [12,13]) is another varie ty ofhaplogroup U present in India (F igure 2) . Unlike theIndian U2, i t has been sampled, a lbei t rare ly, in southernEurop e, th e Near Eas t [12,13] and (according to HVR Isequ ence ident if ica t ion only) a lso in Centra l Asia [18]. Wecalcula ted the co alescence age of this subclus ter in Indiaas 32,000 + 5,500 years: still de ep in late P leist oce ne b utcons ide rab ly younger than tha t fo r U2 . Tab le 2 com paresthe f requ ency of varie t ies of haplogroup U in India , in theTrans -Caucasus popula t ions and in Europe .Typical western-Euras ian mtDNA l ineages found in Indiabelong to haplogroups H, I, J , T, X and to subclusters U1,

Table 1MtDN A haplogroup frequencies ( ) among some Indian andEurasian populat ions.

Population or group of populations

India Wes tern and eastern EurasiaHaplogro up 1 2 3 4 5 6 7affiliationAfrican* 0 0 0 0 0 0 0Eastern 53.3 65.7 60.4 1.5 0.7 61 94.5EurasianWestern 29.3 14.5 20.5 80.9 95.4 30.5 0Eurasian

H 3 1.2 1.8 24.8 41.1 14 0I 2 0 0.7 1,8 2.6 1 0J 0 0.4 0,5 6,7 10.3 2.5 0K 0 0.4 0.2 8.2 4.4 0.5 0T 1 1.7 1.8 11.8 10.1 3~5 0U 23.3 10.3 13.1 21.2 20.8 8 0V 0 0 0 0 3.1 0 0W 0 0.4 2.2 0.9 1.6 1 0X 0 0 0.2 5.5 1.4 0 0

Other s 17.4 19.8 19.1 17.6 3 .9 8.5 5.5The numb ers in italics repres ent the follc~wing popula tions: 1, NorthIndia Uttar Pradesh, n = 1 03, this study); 2, South India AndhraPrades h Telugus, n = 25 0, this study); 3, India total n = 5 50, thisstudy); 4, The Cauc as us - Armenians n = 192, this study), Georgiansn = 138, this study); 5 Europe - Slo vaks n = 129 , this study),

Russians n = 10 0, this study), Czech s n = 95, this study), Estoniansn = 100, this study), Italians n = 99 [27]), Finns n = 49 [16]); 6,Central Asia - Kirghiz n = 95, dedu ced from [18]), Kazakhs n = 55,dedu ced from [18]), Uighurs n = 55, dedu ced from [18]); 7, Tibetn = 54 [26]). *L1 and L2 defined by +3592 Hpal. tLineages that do

not belong te any of the previously established haplogroups.

U4, U5 and K of haplogroup U (Figure 1; Table s 1,2). Fre-quencies o f these l ineages in Indian populat ions are morethan an order of magnitud e lower than in Euro pe: 5.2versus 70 , respect ively (normalised from Table 1). Thi sfinding might be explained by gene f low, as sugges ted pre-viously [2] . Nevertheless , we note that the frequency ofthese mtDNA haplogroups reveals nei ther a s t rongnorth-south, nor language-based gradient : they are foundboth among Hin di speakers from Uttar Pradesh (6 ) andDravidians of Andhra Pradesh (4 ) . Assuming that they arelargely of western-Euras ian origin, we m ay ask when theirspread in India s tarted. To assign a tenta t ive date for theirintroduct ion, we calcula ted the averaged minimal dis tanceof the cor re sponding m tDNA hyperva r iab le reg ionsequences in Indians from the branches shared withwestern Euras ians . We obtained a value for the s ta t is t ic p(see Materials and methods) equal to 0.46, consistent with a


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Brief Communication 1333

Figure 2

23

U72 6A[

3 ~ 168u

. 2e

UlK Cu r r e n t B io lo g y

Reconstruction of haplog roup U lineages found in India. Green boldlines, the backgro un~ of previously characterized haplogroup Ulineages from western Eurasia; red lines, lineage s and hapletype sfound only in India; pink nodes, Dravidic speakers; blue nodes, Hindispeakers. The HVR I mutations at given nucleotide posit ions co mparedwith the Cambridge Reference Sequence [28] are shown less the16,000 prefix near the lines connecting the nodes. Only transversionsare specif ied (for example, 318AT defines an A to T transversion at np16,318). The ancestral node of haplogroup U, marked with an asterisk,differs from the reference sequence by transit ions at nps 0007 3+AIw441) , 7028 +AIM), 12308 +Hinfl) , 11467 -Trul).

divergence time of 9,300_+3,000 years BP. This is anaverage over an unk nown number of various founders and,therefore, does not tell us whether there were one or manymigration waves, or whether there was a continuous long-lasting gradual admixture. Their low frequency but stillgeneral spread all over India plus the estimated time scale,does not support a recent massive Indo-Aryan invasion, atleast as far as maternally inherited genetic lineages are con-cerned. We note, however, that within an error margin thistime estimate is consistent with the arrival to India ofcereals domesticated in the Fertile Crescent [4,19[. Fur-thermore, the spread of these western-Eurasian-specificmtDNA clusters also among Dravidic-speaking p.opulationsof India lends credence to the suggested linguistic connec-tion between Elamite and Dravidic populations [20].Thus, we have shown that the overwhelming majority ofthe so-called western-Eurasian-specific mtD NA lineages inIndian populations, estimated here to be carried by morethan a hundred million contemporary Indians, belong infact to an Indian-specific variety of haplogroup U of a latePleistocene origin. The latter exhibits a direct common

Table 2Freque ncies ( ) of subclusters of haplogroup U in India and insome western-Eurasian populat ions,

Population or group of populations

Indians Armenians , Estonians,Subciuster Georgians Russians, SlovaksU1 2.3 14.4 1.2U2i 77.9 1.0 NFU2e NF 5.2 10.6U3 NF 15.5 4.7U4 4.7 18.6 20.0U5 1.2 11.3 45.9U6 NF NF NFU7 12.7 4.1 NFK 1.2 28.9 12.9Othe r U NF 1.0 4.7Population sizes and their absolute U frequenc y as in Table 1.Subclusters of haplo group U are defined as in [12,13]. U2i andindicate Indian and western-Eurasian variet ies of subcluster U2,respectively (see Figure 2). NF, not found.

U2e

phylogenetic origin with its sister groups found in westernEurasia (Figure 1), but it should not be interpreted in termsof a recent admixture of western Caucasoids with Indianscaused by a puta tive Indo-A ryan invasion 3,000--4,000 yearsBP. From the deep time depth of the split between thepredominant Indian and European haplogroup U varieties,it could be speculated that haplogroup U arose in neither ofthe two regions. This split could have already happened inAfrica, for example, in Ethiopia, where haplogroup U wasrecently described [21].Although there is no strong evidence yet for the presenceof anatomically modern humans in India before35,000-40,000 years ago [22], the earliest estimates of thepresence of modern hum ans in Australia [23] make it verylikely that the subcontinent served as a pathway for east-ward migration of modern humans somewhat earlier andthat it could have been inhabited by them en route as sug-gested by the Souther n Route hypothesis [24,25]. Ourcoalescence age estimate for the mtDNA sub-cluster U2overlaps not only with the corresponding value for theEuropean U5, but with the suggested coalescence age ofthe Indian-specific subset of the predominantly Asianhaplogroup M lineages as well (M.J.B., T.K., W.S.W.,M.E.D., B.B. Rao, J.M. Naidu, e t a / . unpublished observa-tions). Taken together, these data suggest that a commondenominator--most likely beneficial climate condi-ti on s- l ed to the expansion of populations all overEurasia, including t he ancestors of those who now encom-pass most of the mtDNA genome pool of the extant


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1 3 3 4 C u r r e n t B i o l o g y V o l 9 N o 2 2

In d i an s . F u r th e rm o re , t h i s sp ec if i c d i s t r i b u t io n o f m tD N Av ar i e t i e s i n In d i a co m p ared w i th t h e d i s t r i b u t io n o b se rv eda m o n g M o n g o lo i d s a n d t h e C a u c a s oi d p o p u l a t i on s o fw es t e rn E u ras i a F ig u re 1 ) i s, a t p r e sen t , b es t ex p l a in ed b ytw o sep a ra t e l a t e P l e i s to cen e m ig ra t i o n s o f m o d er n h u m an sto In d ia . O n e o f t h em , p o ss ib ly a r r i vin g b y t h e so u th e rnro u te , b ro u g h t t o In d i a an an ces t r a l p o p u la t i o n ca r ry in g h ap -lo g ro u p M an d w as sp read fu r th e r eas tw ard . T h e seco n dm ig ra t i o n b ro u g h t t h e an ces to r s o f h ap lo g ro u p U . A l th o u g ht h e a d m i x t u r e o f th e s e m a j o r w a v e s s t ar t e d p e r h a p s v e r ye a r l y - - e x p l a i n in g t h e s p r e ad o f th e s e m a j o r m t D N A v a ri -e t i e s al l o v e r th e s u b c o n t i n e n t - - i t i s l i k e ly t h at i t h a p -p en ed a f t e r t h e ca r ri e r s o f h ap lo g ro u p M fo u n d th e i r w ayfu r th e r eas t , ex p l a in in g t h e a b sen ce o f h ap lo g ro u p U l i n -eag es am o n g M o n g o lo id p o p u la t i o n s s tu d i ed so fa r.M a t e r i a ls a n d m e t h o d sS a m p l e s f r o m 8 6 L a m b a d i , 6 2 L o 6 a n a ( L a m a n i s p e a k e r s ; I n d o -A r y a nl a n g u a g e s ) , 1 2 T h a ru a n d 1 8 B u k s a ( I n d o -A ry a n l a n g u a g e s ) , 1 2 2 p re -d o m i n a n t l y I n d o -A r y a n l a n g u a g e s p e a k e r s f r o m U t t a r P r a d e s h( G e n B a n k a c c es s i on n u m b e r s A J 2 3 4 9 0 2 - A J 2 3 5 2 0 1 ) a n d a se t o f2 5 0 T e l u g u s a m p l e s ( D r a v i d i c s p ea k e r s ) w e r e s e q u e n c e d f o r h y p e r -v a r i a b le r e g i o n I o f m t D N A a n d t y p e d f o r t h e p r e s e n c e o f m a j o r c o n ti -n e n t - s p e c i f i c m a rk e rs , d e s c r i b e d i n [ 1 1 , 1 6 , 2 6 ] . T h e H V R I p o l y m o rp h i cs i t e s o f a l l 5 5 0 I n d i a n m t D N A s s e q u e n c e d b y u s a re p ro v i d e d i n t h eS u p p l e m e n t a ry m a t e r i a l. ~ F h e p h y l o g e n e t i c a n a l y s i s a l s o i n c l u d e d 1 0 1p u b l is h e d H V R I s e q u e n c e s f r o m s o u t h - w e s t e r n I n d ia [ 6 ] .P h y i o g e n e t i c a n a ly s i s w a s p e r f o r m e d b y r e d u c e d m e d i a n n e t w o r k s[ 1 0 ] , a p p l i e d h e re u s i n g p a rs i m o n y a n a l y s i s o f t h e d a t a . T h e m e d i a nn e t w o rk a n a l y s i s a l l o w s o n e t o r e v e a l s i m u l t a n e o u s l y m u l t i p l e p a ra l l e l ,e q u a l l y p ro b a b l e , p h y l o g e n e t i c p a t h w a y s i n t h e f o rm o f r e t i c u l a t i o n si n d u c e d b y h i g h l y v a r i a b l e m a rk e rs . T h e d i s t i n c t m t D N A l i n e a g e c lu s -t e r s a re re f e r re d t o h e re a s h a p l o g ro u p s . T h e t i m e t o t h e m o s t r e c e n tc o m m o n a n c e s t o r o f a c l u s t e r o f l i n e a g e s ( h a p l o g r o u p ) o r , w h e r ea p p ro p r i a t e , a s u b -c l u s t e r i n s i d e a p a r t i c u l a r h a p l o g ro u p , w a s c a l c u -l a t e d a s d e s c r i b e d [ 1 7 ] , u s i n g a n e s t i m a t o r p , w h i c h i s t h e a v e ra g et r a n s it i o n al d i s t a n c e f r o m t h e f o u n d e r h a p l o t y p e s e q u e n c e .S u p p l e m e n t a r y m a t e r i a lS u p p l e m e n t a ry m a t e r i a l i n c l u d i n g a t a b l e l is t i n g t h e m t D N A H V R Is e q u e n c e p o l y m o r p h i s m s i n d i f f er e n t I n d ia n p o p u l a t i o n s a n d a m o r ed e t a i l e d d e s c r i p t i o n o f t h e m a t e r i a l s a n d m e t h o d s i s a v a i l a b le a th t t p : / / c u r r e n t - b i o l o g y . c o m / s u p m a t / s u p m a t i n . h t m .

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bogus claims about aryan invasion

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