RAMAYANA NOT A WORK OF FICTION
New research carried out by anthropological scientists from the Estonian Biocentre and the University of Delhi claims that events of the mythological epic Ramayana occurred in reality thousands of years ago.
Hyderabad: New research carried out by anthropological scientists from the Estonian Biocentre and the University of Delhi claims that events of the mythological epic Ramayana occurred in reality thousands of years ago. Scientists say that results of their genetic studies, with existing data, show genetic signatures of tribal groups featured in the Ramayana such as the Gonds, Kols and Bhils. Gonds are a prominent group in Adilabad district of Telangana.
Researchers claimed that populations in the Indian subcontinent can trace their ancestors to more than 60,000 years back. Scientists say that this is proof of the authenticity and actual occurrence of the events described in Ramayana, which would have occurred more than 12,000 years ago. The Gonds, Kols and Bhils are believed to be the ancient tribal groups of the region and have found mention in the Ramayana. Authenticity of the mythological text has been questioned several times. While there have been voices proclaiming the authenticity of the Ramayana, research to prove it has increased in recent times.
Dr Vadlamudi Raghavendra Rao, professor of anthropology, University of Delhi, and one of the authors of the study, said, “Definitely, the events described in Ramayana occurred in real. Our research has showed close genetic affinity of these tribes to other ethnic groups. We have shown that there is continuity in the populations groups living here. Other researchers are working to prove other angles of this.”
The study was carried out by Estonian Biocentre researcher Gyaneshwar Chaubey, Institute of Scientific Research on Vedas, Dr Saroj Bala and Dr Raghavendra Rao. The Bhil, Kol and Gond are three major Indian tribes that have been widely acknowledged in the epic Ramayana, particularly in the chapters Ayodhyakanda, Aranyakanda and Kishkindhakanda. Gonds are prominently found in Adilabad district and other states and number about 40 lakh.
The research study says since these tribes are inhabitants of the country since the Stone Age, their genetic affinity to existing populations show the authenticity of the Ramayana.
Researchers said these tribal groups form a closed cluster with Dravidian groups, known as inhabitants of South India.
http://www.deccanchronicle.com/150618/nation-current-affairs/article/ramayana-real-say-experts
RAMAYANA NOT A WORK OF FICTION
10/26/2015 PLOS ONE: Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic Ramayana
http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0127655 1/8
Correction
24 Jul 201 5: The PLOS ONE Staff (201 5) Correction: Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic Ramayana.
PLoS ONE 1 0(7): e01 34200. doi: 1 0.1 371 /journal.pone.01 34200 | View correction
Abstract
Kol, Bhil and Gond are some of the ancient tribal populations known from the Ramayana, one of the Great epics of India. Though there have been studies about their affinity based on classical and haploid genetic markers, the molecular insights of their
relationship with other tribal and caste populations of extant India is expected to give more clarity about the the question of continuity vs. discontinuity.
Citation: Chaubey G, Kadian A, Bala S, Rao VR (201 5) Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic
Ramayana. PLoS ONE 1 0(6): e01 27655. doi:1 0.1 371 /journal.pone.01 27655
Academic Editor: Kumarasamy Thangaraj, Centre for Cellular and Molecular Biology, INDIA
Received: September 7, 201 4; Accepted: April 1 7, 201 5; Published: June 1 0, 201 5
Copyright: © 201 5 Chaubey et al. This is an open access article distributed under the terms of the Creative Commons
Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author
and source are credited
Data Availability: Data have been deposited to Figshare: http://dx.doi.org/1 0.6084/m9.figshare.1 391 920. The data are also
available at the data repository of the Estonian Biocentre: www.ebc.ee/free_data.
Funding: The authors received no specific funding for this work. GC is supported by Center of Excellence of Estonian
Biocentre and Institutional Grant IUT241 .
Competing interests: Coauthor Dr. Gyaneshwer Chaubey is a PLOS ONE Editorial Board member. This does not alter the
authors' adherence to all the PLOS ONE policies on sharing data and materials.
Our survey on mythological sources has revealed detailed information about the ancient Indian society structure as well as relations of different tribal and caste groups and their rituals [1 0,11 ,1 6–1 8].
The Bhils are primarily from Central India and speak the Bhil language [28]. They have significant presence in states of Gujarat, Madhya Pradesh, Chhattisgarh, Maharashtra and Rajasthan as well as in the northeastern state of Tripura.
Published: June 1 0, 201 5 DOI: 1 0.1 371 /journal.pone.01 27655
Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic
Ramayana
Gyaneshwer Chaubey , Anurag Kadian, Saroj Bala, Vadlamudi Raghavendra Rao
10/26/2015 PLOS ONE: Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic Ramayana
http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0127655 2/8
Uttar Pradesh.
More than 25 years of genetic research on Indian tribal and caste populations involving classical markers to mtDNA/Y chromosome
and more recently autosomes, have indicated complex demographic history of the subcontinent [3,30–39].
Therefore, in the present study, we make an attempt to evaluate two schools of thought emerging from the current scenario.
Material and Methods
This study was performed using control samples collected, genotyped and published for various population studies conducted in the last few years (S1 Table) [37–39,46,61 –63].
We performed PC analysis using smartpca programme (with default settings) of the EIGENSOFT package [67] in order to capture genetic variability described by the first 5 components. The fraction of the total variation described by a PC is the ratio of its
eigenvalue to the sum of all eigenvalues.
Mean pairwise differences between different population groups were computed using Fst distance measure by following the methods as described by Cockerham and Weir [69], Phylip [70] and MEGA [71 ] were used to construct the tree. The Plink software
[66] was used to calculate the genetic diversity and to find the 25 nearestneighbours for the Bhil, Kol and Gond individuals.
Results and Discussion
We combined hundreds of thousands of autosomal markers generated from different studies (S1 Table) [37–39,46,61 –63] and specifically looked into the population structure of Indian groups mentioned in classical literature.
10/26/2015 PLOS ONE: Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic Ramayana
http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0127655 3/8
Taken together, these results support the second hypothesis suggesting that all Indians, irrespective of their caste or tribal affiliations, share a common genetic ancestry, which is undoubtedly founded over the indigenous ASI component.
Our extended analysis on Fst, PCA and ADMIXTURE showed similar genome composition of these tribal populations, carrying both the ANE and ASI components (Fig 1 a–1 c, Table 1 and S1 Fig). We also calculated the genetic diversity of these populations with their neighbours (Table 2).
10/26/2015 PLOS ONE: Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic Ramayana
http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0127655 4/8
Table 1. The ANE and ASI admixture proportions of Bhil, Kol and Gond with respect to other South Asian groups.
doi:1 0.1 371 /journal.pone.01 27655.t001
Table 2. The genetic diversity calculations of Bhil, Kol and Gond with respect to other South Asian groups.
doi:1 0.1 371 /journal.pone.01 27655.t002
The haplotype based fineSTRUCTURE [72] analysis showed that the studied populations (Bhil, Kol and Gond) received nearly all of their chunks from the Indian closeby populations (Fig 2a and S3 Fig).
Supporting Information
S1 Fig. Neighbour Joining (NJ) tree world populations inferred from Fst distances of genomewide data.
In the inset, the heatmap showing the inter and intra regional genetic affinity of the three tribal populations under investigation.
doi:1 0.1 371 /journal.pone.01 27655.s001
(TIF)
S2 Fig. The plot of shared drift obtained by the f3 = (Yoruba; Bhil/Kol/Gond, X).
The f3 values are plotted on Y axis against the X targeted populations on X axis. C_Asia Central Asia, IN_IE_Caste Indian IndoEuropean Caste, IN_IE_Tribe—Indian IndoEuropean Tribe, IN_DRA_Caste Indian Dravidian Caste, IN_DRA_Tribe—Indian
Dravidian Tribe, IN_AA Indian Austroasiatic (Munda).
doi:1 0.1 371 /journal.pone.01 27655.s002
(TIF)
S3 Fig. Coancestry matrix plotted from fineSTRUTCURE analysis, showing the chunks donated by other Eurasian populations to the Bhil, Kol and
Gond populations.
doi:1 0.1 371 /journal.pone.01 27655.s003
(TIF)
S4 Fig. Placement of Kol, Bhil and Gond individuals over the 37 clades obtained from the fineSTRUCTURE analysis.
doi:1 0.1 371 /journal.pone.01 27655.s004
(TIF)
S1 Table. The details of the populations (number of individuals and number of SNPs, used in the present study.
doi:1 0.1 371 /journal.pone.01 27655.s005
(XLSX)
10/26/2015 PLOS ONE: Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic Ramayana
http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0127655 5/8
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New research carried out by anthropological scientists from the Estonian Biocentre and the University of Delhi claims that events of the mythological epic Ramayana occurred in reality thousands of years ago.
Hyderabad: New research carried out by anthropological scientists from the Estonian Biocentre and the University of Delhi claims that events of the mythological epic Ramayana occurred in reality thousands of years ago. Scientists say that results of their genetic studies, with existing data, show genetic signatures of tribal groups featured in the Ramayana such as the Gonds, Kols and Bhils. Gonds are a prominent group in Adilabad district of Telangana.
Researchers claimed that populations in the Indian subcontinent can trace their ancestors to more than 60,000 years back. Scientists say that this is proof of the authenticity and actual occurrence of the events described in Ramayana, which would have occurred more than 12,000 years ago. The Gonds, Kols and Bhils are believed to be the ancient tribal groups of the region and have found mention in the Ramayana. Authenticity of the mythological text has been questioned several times. While there have been voices proclaiming the authenticity of the Ramayana, research to prove it has increased in recent times.
Dr Vadlamudi Raghavendra Rao, professor of anthropology, University of Delhi, and one of the authors of the study, said, “Definitely, the events described in Ramayana occurred in real. Our research has showed close genetic affinity of these tribes to other ethnic groups. We have shown that there is continuity in the populations groups living here. Other researchers are working to prove other angles of this.”
The study was carried out by Estonian Biocentre researcher Gyaneshwar Chaubey, Institute of Scientific Research on Vedas, Dr Saroj Bala and Dr Raghavendra Rao. The Bhil, Kol and Gond are three major Indian tribes that have been widely acknowledged in the epic Ramayana, particularly in the chapters Ayodhyakanda, Aranyakanda and Kishkindhakanda. Gonds are prominently found in Adilabad district and other states and number about 40 lakh.
The research study says since these tribes are inhabitants of the country since the Stone Age, their genetic affinity to existing populations show the authenticity of the Ramayana.
Researchers said these tribal groups form a closed cluster with Dravidian groups, known as inhabitants of South India.
http://www.deccanchronicle.com/150618/nation-current-affairs/article/ramayana-real-say-experts
RAMAYANA NOT A WORK OF FICTION
10/26/2015 PLOS ONE: Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic Ramayana
http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0127655 1/8
Correction
24 Jul 201 5: The PLOS ONE Staff (201 5) Correction: Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic Ramayana.
PLoS ONE 1 0(7): e01 34200. doi: 1 0.1 371 /journal.pone.01 34200 | View correction
Abstract
Kol, Bhil and Gond are some of the ancient tribal populations known from the Ramayana, one of the Great epics of India. Though there have been studies about their affinity based on classical and haploid genetic markers, the molecular insights of their
relationship with other tribal and caste populations of extant India is expected to give more clarity about the the question of continuity vs. discontinuity.
In this study, we scanned >97,000 of single nucleotide polymorphisms among three major ancient tribes mentioned in Ramayana, namely Bhil, Kol and Gond.
The results obtained were then compared at inter and intra population levels with neighboring and other world populations.
Using various statistical methods, our analysis suggested that the genetic architecture of these tribes (Kol and Gond) was largely similar to their surrounding tribal and caste populations, while Bhil showed closer affinity with Dravidian and Austroasiatic (Munda) speaking tribes.
The haplotype based analysis revealed a massive amount of genome sharing among Bhil, Kol, Gond and with other ethnic groups of South Asian descent.
On the basis of genetic component sharing among different populations, we anticipate their primary founding over the indigenous Ancestral South Indian (ASI) component has prevailed in the genepool over the last several thousand years.
Citation: Chaubey G, Kadian A, Bala S, Rao VR (201 5) Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic
Ramayana. PLoS ONE 1 0(6): e01 27655. doi:1 0.1 371 /journal.pone.01 27655
Academic Editor: Kumarasamy Thangaraj, Centre for Cellular and Molecular Biology, INDIA
Received: September 7, 201 4; Accepted: April 1 7, 201 5; Published: June 1 0, 201 5
Copyright: © 201 5 Chaubey et al. This is an open access article distributed under the terms of the Creative Commons
Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author
and source are credited
Data Availability: Data have been deposited to Figshare: http://dx.doi.org/1 0.6084/m9.figshare.1 391 920. The data are also
available at the data repository of the Estonian Biocentre: www.ebc.ee/free_data.
Funding: The authors received no specific funding for this work. GC is supported by Center of Excellence of Estonian
Biocentre and Institutional Grant IUT241 .
Competing interests: Coauthor Dr. Gyaneshwer Chaubey is a PLOS ONE Editorial Board member. This does not alter the
authors' adherence to all the PLOS ONE policies on sharing data and materials.
Introduction
Knowledge about the past comes through different disciplines where researchers look at history through different lenses. And in many cases, these interdisciplinary studies land on the same conclusions [1 ,2].
Knowledge about the past comes through different disciplines where researchers look at history through different lenses. And in many cases, these interdisciplinary studies land on the same conclusions [1 ,2].
However, in case of India, investigations from
different disciplines have historically been highly contrasting [3,4].
different disciplines have historically been highly contrasting [3,4].
India, also known as a ‘land of spiritual heritage’, has a deep history of civilisation, which is embedded in to multiple oral, traditional and written records. Much of this knowledge is rooted in oldest scriptures, the Vedas, which are four in number, namely Rigveda, Yajurveda, Samaveda and Atharvaveda.
Then, there are Puranas, Upanishads, Brahmanas and Aranyakas, of which Vedas are said to be the precursors [5].
There is no consensus among historians regarding the date of compilation of the Vedas as well as the historical dates for the various Puranas, Upanishads and epics [6–1 0].
A comperative analysis of such mythological sources may provide a concencus about the structuring of the ancient societies and rituals.
More recently, some scholars have provided strong evidence about the chronology of these events hinting at a deep rooted civilization, developing indigenously for over several thousand years [8,11 –1 5].
Our survey on mythological sources has revealed detailed information about the ancient Indian society structure as well as relations of different tribal and caste groups and their rituals [1 0,11 ,1 6–1 8].
In many of these literary sources, names of various castes and tribal groups have been mentioned, including those of several surviving tribal groups (e.g. Bhil or Bheel, Kol, Gond, Savara, Oraon,
Kirata, Ahirs, Nagas etc) [1 7–23]. It is already evident that during the Ramayana era, Indian society was well stratified [1 6,1 7,21 ,23–26].
Kirata, Ahirs, Nagas etc) [1 7–23]. It is already evident that during the Ramayana era, Indian society was well stratified [1 6,1 7,21 ,23–26].
The Bhil, Kol and Gond are three major Indian tribes that have been widely acknowledged in the epic Ramayana, particularly in the portions known as the Ayodhyakanda, Aranyakanda and Kishkindhakanda [1 9,20,22–27].
It should be emphasised here that, Gond and Bhil are the top two tribal populations of modern India in terms of population size [28].
The Bhils are primarily from Central India and speak the Bhil language [28]. They have significant presence in states of Gujarat, Madhya Pradesh, Chhattisgarh, Maharashtra and Rajasthan as well as in the northeastern state of Tripura.
Bhils are further divided into a number of endogamous territorial divisions, which in turn have a number of clans and lineages [22].
The Kol tribe in Uttar Pradesh is found mainly in the districts of Mirzapur, Varanasi, Banda and Allahabad [28]. It is the largest tribe found in the state
Published: June 1 0, 201 5 DOI: 1 0.1 371 /journal.pone.01 27655
Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic
Ramayana
Gyaneshwer Chaubey , Anurag Kadian, Saroj Bala, Vadlamudi Raghavendra Rao
10/26/2015 PLOS ONE: Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic Ramayana
http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0127655 2/8
Uttar Pradesh.
They are said to have migrated from Central India some five centuries ago [28]. The Kol are further divided into a number of exogamous clans, such as the Rojaboria, Rautia, Thakuria, Monasi, Chero and Barawire.
The Gond people are spread over the states of Madhya Pradesh, eastern Maharashtra (Vidarbha), Chhattisgarh, Uttar Pradesh and Telangana [28]. With over four million people, they are the largest tribe in Central India. They speak the Gondi language, which is related to present Dravidian
language family [28,29].
language family [28,29].
More than 25 years of genetic research on Indian tribal and caste populations involving classical markers to mtDNA/Y chromosome
and more recently autosomes, have indicated complex demographic history of the subcontinent [3,30–39].
Alongwith debate over initial peopling of the subcontinent, the major hot topic now shifted towards the population expansion and admixture during and after Neolithic times [37–40].
However, large number of individuals as well as genetic markers are required to reach any firm
conclusions. Thus, the strict endogamy and social structure make South Asia much more complex, unlike to Europe, where genetic
analysis of a population can predict the genetic structure of immediate neighbor with some confidence.
conclusions. Thus, the strict endogamy and social structure make South Asia much more complex, unlike to Europe, where genetic
analysis of a population can predict the genetic structure of immediate neighbor with some confidence.
In recent years, there has been an increase in the number of indepth genetic studies focussing on the genetic structure of the populations of India
[35,37,40–48], but none of them have related specific tribal populations mentioned in the traditional literatures.
[35,37,40–48], but none of them have related specific tribal populations mentioned in the traditional literatures.
Therefore, in the present study, we make an attempt to evaluate two schools of thought emerging from the current scenario.
The first school suggests that the tribal people are the aboriginal inhabitants, while the later migrants, i.e., the Dravidians followed by the Aryans have pushed them back in to small pockets in South India [49–52].
According to this school, the caste system was
established by the aforementioned later migrants [11 ,50,52,53].
established by the aforementioned later migrants [11 ,50,52,53].
The alternative hypothesis advocates that all the caste and tribal populations of India have Paleolithic roots and share a common origin [3,1 5,33,54–60].
The differentiation observed in modern
South Asian populations is mainly derived by strict endogamy, long term isolation and several evolutionary forces.
South Asian populations is mainly derived by strict endogamy, long term isolation and several evolutionary forces.
More specifically, relying on each other, first, we seek to investigate the continuity vs. discontinuity of the genetic thread connecting the different
populations of India.
populations of India.
Second, keeping in mind the pivotal information extracted from Ramayana, we look specifically into the question: whether and to what extent the three major tribes (Bhil, Kol and Gond) share their genetic ancestry among them as well as with other contemporary caste and tribal populations?
Material and Methods
This study was performed using control samples collected, genotyped and published for various population studies conducted in the last few years (S1 Table) [37–39,46,61 –63].
All the ethical guideline have been followed. The tribal and caste populations grouped according to their language group.
We grouped populations in to “Transitional” who have known information of language change in recent time [64,65]. A check for closely related individuals was carried out within each population study by calculating average identity by state (IBS) scores for all pairs of individuals [66].
We used PLINK 1 .07 [66] in order to filter our dataset to include only SNPs on the 22 autosomal chromosomes with minor allele frequency >1 % and genotyping success >99%.
As background linkage disequilibrium (LD) can affect both PCA [67] and ADMIXTURE [68], we thinned the dataset by removing one SNP of any pair, in strong LD r2>0.4, in a window of 200 SNPs (sliding the window by 25 SNPs at a time).
We performed PC analysis using smartpca programme (with default settings) of the EIGENSOFT package [67] in order to capture genetic variability described by the first 5 components. The fraction of the total variation described by a PC is the ratio of its
eigenvalue to the sum of all eigenvalues.
In the final settings, we ran Admixture with a random seed number generator on the LDpruned dataset twentyfive times at K = 2 to K = 1 2. Since the top values of the resulting loglikelihood scores were stable (virtually identical) within the runs of each K from K = 2 to K = 1 0, we can claim that convergence at global maximum was achieved. Thus,
we omitted runs at K = 11 to K = 1 2 from further analysis.
we omitted runs at K = 11 to K = 1 2 from further analysis.
Mean pairwise differences between different population groups were computed using Fst distance measure by following the methods as described by Cockerham and Weir [69], Phylip [70] and MEGA [71 ] were used to construct the tree. The Plink software
[66] was used to calculate the genetic diversity and to find the 25 nearestneighbours for the Bhil, Kol and Gond individuals.
To investigate the derived allele sharing of Bhil, Kol and Gond with the Eurasian populations, we computed f3 statistics [37], taking African as an outgroup.
For haplotypebased analysis (fineSTRUCTURE) [72], we made two different runs—first by taking all the
Eurasian populations and second exclusively on the Central Asian, Pakistani and Indian populations.
Eurasian populations and second exclusively on the Central Asian, Pakistani and Indian populations.
For the fine STRUCTURE analysis, first samples were phased with Beagle 3.3.2 [73]. A coancestry matrix was constructed using ChromoPainter [72],
fineSTRUCTURE was used to perform an MCMC iteration using 1 0000000 burning runtime and 1 0000 MCMC samples.
fineSTRUCTURE was used to perform an MCMC iteration using 1 0000000 burning runtime and 1 0000 MCMC samples.
A tree was built using fineSTRUCTURE with the default settings. All these information are plotted for the Bhil, Kol and Gond as a recipient of
number of chunks from one another as well as from other ethnic group.
number of chunks from one another as well as from other ethnic group.
Results and Discussion
We combined hundreds of thousands of autosomal markers generated from different studies (S1 Table) [37–39,46,61 –63] and specifically looked into the population structure of Indian groups mentioned in classical literature.
To find out the population clustering, we first ran the Fst (population differentiation) algorithm [69] and drew a tree [70,71 ], rooting out the African populations (S1 Fig).
All the Indian populations, except the present TibetoBurman speaking populations, are well separated from other continental populations and form a major cluster comprising present populations speaking IndoEuropean, Dravidian and Austroasiatic (Munda) languages (S1 Fig).
The Pakistani populations are scattered in different clusters, where few of them (Sindhi, Pathan and Burusho) cluster loosely with Indians; Hazaras show an affinity toward Central Asians, and Balochi, Brahui and Makrani confirm an intermediate position because of shared recent African ancestry and gene flow [38,74,75].
The Bhil, Kol and Gond showed a closer affinity among them as well as with the extent IndoEuropean, Transitional and Munda speaking populations (Fig 1 a and S1 Fig).
10/26/2015 PLOS ONE: Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic Ramayana
http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0127655 3/8
Fig 1.
a) Regionwise population differentiation (Fst) analysis of Bhil, Kol Gond with the Indian and other regional populations
a) Regionwise population differentiation (Fst) analysis of Bhil, Kol Gond with the Indian and other regional populations
b) PCA (Principle Component Analysis) of Eurasian populations showing the placement of Bhil, Kol and Gond populations over the South Asian cline; the mean value of populationwise clustering of Bhil, Kol and Gond is zoomedin inset figure. IE IndoEuropean, DRA Dravidian
c) Individualwise ancestry proportion analysis inferred from ADMIXTURE representing ten
ancestral populations of the world (K = 1 0).
doi:1 0.1 371 /journal.pone.01 27655.g001
ancestral populations of the world (K = 1 0).
doi:1 0.1 371 /journal.pone.01 27655.g001
To get more deeper insight, we have used PCA (principle component analysis)[67] and ADMIXTURE [68], analysis using the same parameters as in our previous studies [38,39,45].
These analyses strengthened the inferences drawn from the Fst analysis. The PCA on Eurasians placed Indian populations between East and West Eurasia (Fig 2a).
The cline of Indian subcontinent ranges
from Pakistani populations (closer to West Eurasians) to Indian Munda groups (closer to East Eurasians). Departing from its geographical position, Bhil was clustering together with Scheduled castes and Scheduled tribe populations of Uttar Pradesh
(Harijan), Andhra Pradesh (Kamsali) and Karnataka (North Kannadi) states. Kol is joined with the neighbouring populations alongwith the Indiancline, while Gond was deflating away from the Indian cline by uniting with the Munda speakers (Fig 1 b).
from Pakistani populations (closer to West Eurasians) to Indian Munda groups (closer to East Eurasians). Departing from its geographical position, Bhil was clustering together with Scheduled castes and Scheduled tribe populations of Uttar Pradesh
(Harijan), Andhra Pradesh (Kamsali) and Karnataka (North Kannadi) states. Kol is joined with the neighbouring populations alongwith the Indiancline, while Gond was deflating away from the Indian cline by uniting with the Munda speakers (Fig 1 b).
Further, we assessed the proportion of individualwise ancestry drawn from a given number of inferred populations (K) using a maximum likelihood based approach implemented in ADMIXTURE.
Fig 2.
a) The number of chunks donated at inter and intra populations level for the Bhil, Kol and Gond with respect to the Indian, Central Asian and the Pakistani populations. b) Plot of 25 nearest neighbors of Bhil, Kol and Gond individuals. The match
population individuals are colored in grey.
doi:1 0.1 371 /journal.pone.01 27655.g002
Consistent with previous observations [37,38], the South Asian populations’ genome are mainly madeup of two major components, which are distributed across the length and breadth of the subcontinent (Fig 1 c).
a) The number of chunks donated at inter and intra populations level for the Bhil, Kol and Gond with respect to the Indian, Central Asian and the Pakistani populations. b) Plot of 25 nearest neighbors of Bhil, Kol and Gond individuals. The match
population individuals are colored in grey.
doi:1 0.1 371 /journal.pone.01 27655.g002
Consistent with previous observations [37,38], the South Asian populations’ genome are mainly madeup of two major components, which are distributed across the length and breadth of the subcontinent (Fig 1 c).
Alongn with these two major components, there are
four minor componets over the periphery of the subcontinent—the European and the Middle eastern components can be seen in Pakistani and northwest Indian populations, whilest the East/Southeast Asian components are present in nearby Munda and TibetoBurman speakers.
four minor componets over the periphery of the subcontinent—the European and the Middle eastern components can be seen in Pakistani and northwest Indian populations, whilest the East/Southeast Asian components are present in nearby Munda and TibetoBurman speakers.
(Fig 1 c). The geographical distribution of the dark green component (ASI or Ancestral South Indian unique to the subcontinent) was largely limited to the Indian subcontinent, and can be seen among all the populations of the subcontinent albeit in variable amount, whereas the second major component (light green: ANI or Ancestral North Indian (now ANE Ancestral North Eurasian [76])) was shared with Central Asia, the Caucasus, Middle East and Europe (Fig 1 c).
The geographical origin of light green component (ANI or ANE) is so far unclear and more research is needed from unsampled area as well as from ancient DNA; however, the time of spread of this component from its origin place (either of any; the Caucasus, Near East, Indus Valley, or Central Asia) has happened more than 1 2.5 thousand years before [38], which is significantly earlier than the purported expansion of Dravidians and Aryans languages from outside the subcontinent.
Notably, the Andaman Islanders are not the only population carrying the ASI component exclusively, as was suggested before [37].
Austroasiatic speakers (more precisely, the South Munda) of the subcontinent also seem to possess the ASI component in near unadulterated form (Fig 1 c).
More research with complete genome analysis would be required to clear the geographic center of the ANE component; however, it is evident from the present
analysis that the dark green component (ASI) can be considered as a connecting thread for all the Indian populations (Fig 1 c).
analysis that the dark green component (ASI) can be considered as a connecting thread for all the Indian populations (Fig 1 c).
Taken together, these results support the second hypothesis suggesting that all Indians, irrespective of their caste or tribal affiliations, share a common genetic ancestry, which is undoubtedly founded over the indigenous ASI component.
Our second question revolved around the three tribal populations mentioned in the ancient epic, their genome composition and affiliation with the surrounding caste and tribal populations. Based on information from Ramayana, we have considered these tribal populations to be ancient inhabitants of India, surviving from the times of the Stone Age [1 9,23].
If we assume that their genome carry the signature of peopling of ancient time, the assessment of their genomes and comparison with modern populations would test the scenario of continuity vs. discontinuity of prehistoric heritage.
In case of continuity, we should see largely similar genome composition among contemporary caste and tribal populations of modern India. On the other hand, in case of discontinuity, these tribal populations should show a unique genome composition or they should emerge as an outliers in our cluster based analysis.
Our extended analysis on Fst, PCA and ADMIXTURE showed similar genome composition of these tribal populations, carrying both the ANE and ASI components (Fig 1 a–1 c, Table 1 and S1 Fig). We also calculated the genetic diversity of these populations with their neighbours (Table 2).
The diversity of Kol, Bhil and Gond didn’t show any significant deviation from their neighbouring extent
IndoEuropean, Dravidian and Munda speaking groups.
IndoEuropean, Dravidian and Munda speaking groups.
10/26/2015 PLOS ONE: Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic Ramayana
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Table 1. The ANE and ASI admixture proportions of Bhil, Kol and Gond with respect to other South Asian groups.
doi:1 0.1 371 /journal.pone.01 27655.t001
Table 2. The genetic diversity calculations of Bhil, Kol and Gond with respect to other South Asian groups.
doi:1 0.1 371 /journal.pone.01 27655.t002
The shared drift statistics analysis (f3) suggested that most of the derived alleles of Bhil, Kol and Gond are overwhelmingly shared with Indian caste and tribal populations (S2 Fig).
Gond, Dravidian tribes and Austroasiatic (Munda) groups shared the highest derived allele with Bhils.
IndoEuropean castes, Gond and Dravidian tribes were closest with Kol.
Whereas, Munda, transitional and Dravidian tribal groups shared the peak derived alleles with Gonds.
The haplotype based fineSTRUCTURE [72] analysis showed that the studied populations (Bhil, Kol and Gond) received nearly all of their chunks from the Indian closeby populations (Fig 2a and S3 Fig).
Leaving out the number of chunks coming from the same population, chunk donors for Bhil and Kol were coming from all the major Indian ethnic groups, while for the Gond, Indian Transitional and Munda groups were the major chunk donors.
More specifically the haplotype based sharing analysis is in congruent with the f3 statistics. The fineSTRUCTURE clustering analysis revealed 37 clusters when we have included Iranian,
Central Asian, Pakistani, Indian and Cambodian populations (S4 Fig).
Central Asian, Pakistani, Indian and Cambodian populations (S4 Fig).
Most of the Indian populations unite in Indian specific clusters except Kashmiri Pandits and few Gujarati individuals who fell together with the Sindhi and Pathan individuals in Pakistani specific clade.
Our targeted populations are dispersed in various clades. All the Bhil individuals form a tight cluster with the individuals mainly from Dravidian caste, few IndoEuropean and Transitional individuals.
Most of the Kol and Gond individuals show a higher
level of variation by falling in to distinct clusters. To make an individualwise comparison, we plotted top twenty five closest neighbours of studied populations (Fig 2b).
level of variation by falling in to distinct clusters. To make an individualwise comparison, we plotted top twenty five closest neighbours of studied populations (Fig 2b).
It was expected that any population members would be closest to themselves and thereafter to members of other populations, which was also pertinent in the present study. Consistent with the above observations drawn from Fst, PCA, ADMIXTURE, f3 statistics and fineSTRUCTURE, there is no signature of large scale population replacement in the Indian subcontinent.
In conclusion, our high resolution analysis portraying the three ancient tribal populations, strongly rejects any incoming genetic signal of large scale recent (during the postNeolithic) migration either of the present Dravidian or the IndoEuropean speaking populations to the subcontinent.
We also concluded that the Indian populations preserve strong genetic signatures in support of a
common ancestry. The studied tribal populations do share large number of genome among theselves as well as from o caste and tribal populations.
common ancestry. The studied tribal populations do share large number of genome among theselves as well as from o caste and tribal populations.
Notebly, the placement of various populations along the Indian cline is not solely governed by the geography, but also by the castetribe interaction and various other selectional forces.
These patterns point to a complex demographic history of the subcontinent which has been shaped insitu by admixture events at different time scale, as well as by intricate geographical heterogeneity and long term effect of several evolutionary forces.
Supporting Information
S1 Fig. Neighbour Joining (NJ) tree world populations inferred from Fst distances of genomewide data.
In the inset, the heatmap showing the inter and intra regional genetic affinity of the three tribal populations under investigation.
doi:1 0.1 371 /journal.pone.01 27655.s001
(TIF)
S2 Fig. The plot of shared drift obtained by the f3 = (Yoruba; Bhil/Kol/Gond, X).
The f3 values are plotted on Y axis against the X targeted populations on X axis. C_Asia Central Asia, IN_IE_Caste Indian IndoEuropean Caste, IN_IE_Tribe—Indian IndoEuropean Tribe, IN_DRA_Caste Indian Dravidian Caste, IN_DRA_Tribe—Indian
Dravidian Tribe, IN_AA Indian Austroasiatic (Munda).
doi:1 0.1 371 /journal.pone.01 27655.s002
(TIF)
S3 Fig. Coancestry matrix plotted from fineSTRUTCURE analysis, showing the chunks donated by other Eurasian populations to the Bhil, Kol and
Gond populations.
doi:1 0.1 371 /journal.pone.01 27655.s003
(TIF)
S4 Fig. Placement of Kol, Bhil and Gond individuals over the 37 clades obtained from the fineSTRUCTURE analysis.
doi:1 0.1 371 /journal.pone.01 27655.s004
(TIF)
S1 Table. The details of the populations (number of individuals and number of SNPs, used in the present study.
doi:1 0.1 371 /journal.pone.01 27655.s005
(XLSX)
10/26/2015 PLOS ONE: Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic Ramayana
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Author Contributions
Conceived and designed the experiments: GC. Performed the experiments: GC AK SB VRR. Analyzed the data: GC. Contributed
reagents/materials/analysis tools: AK SB VRR. Wrote the paper: GC AK SB VRR.
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Correction
24 Jul 201 5: The PLOS ONE Staff (201 5) Correction: Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic Ramayana.
PLoS ONE 1 0(7): e01 34200. doi: 1 0.1 371 /journal.pone.01 34200 | View correction
Conceived and designed the experiments: GC. Performed the experiments: GC AK SB VRR. Analyzed the data: GC. Contributed
reagents/materials/analysis tools: AK SB VRR. Wrote the paper: GC AK SB VRR.
References
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pmid:1 261 0536 doi: 1 0.1 038/ng1 1 1 3
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Asia. Science 326: 1 541 –1 545. doi: 1 0.1 1 26/science.1 1 77074. pmid:20007900
Chaubey G, Metspalu M, Kivisild T, Villems R (2007) Peopling of South Asia: investigating the castetribe continuum in India. Bioessays 29: 91 –1 00.
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Boivin N (2007) Anthropological, historical, archaeological and genetic perspectives on the origins of caste in South Asia. The Evolution and History of
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Correction
24 Jul 201 5: The PLOS ONE Staff (201 5) Correction: Genetic Affinity of the Bhil, Kol and Gond Mentioned in Epic Ramayana.
PLoS ONE 1 0(7): e01 34200. doi: 1 0.1 371 /journal.pone.01 34200 | View correction
Abstract
Kol, Bhil and Gond are some of the ancient tribal populations known from the Ramayana, one of the Great epics of India.
Kol, Bhil and Gond are some of the ancient tribal populations known from the Ramayana, one of the Great epics of India.
Though there have been studies about their affinity based on classical and haploid genetic markers, the molecular insights of their relationship with other tribal and caste populations of extant India is expected to give more clarity about the the question of continuity vs. discontinuity.
In this study, we scanned >97,000 of single nucleotide polymorphisms among three major ancient tribes mentioned in Ramayana, namely Bhil, Kol and Gond.
The results obtained were then compared at inter and intra population levels with neighboring and other world populations.
Using various statistical methods, our analysis suggested that the genetic architecture of these tribes (Kol and Gond) was largely similar to their surrounding tribal and caste populations, while Bhil showed closer affinity with Dravidian and Austroasiatic (Munda) speaking tribes.
The haplotype based analysis revealed a massive amount of genome sharing among Bhil, Kol, Gond and with other ethnic groups of South Asian descent.
On the basis of genetic component sharing among different populations, we anticipate their primary founding over the indigenous Ancestral South Indian (ASI) component has prevailed in the genepool over the last several thousand years.
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