Genetic Sketch of the Six Population Groups of Rajasthan
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ANTHROPOLOGICAL SCIENCE Vol. 119(3), 259–264, 2011 Genetic sketch of the six population groups of Rajasthan: a study based on 12 autosomal loci Rima DADA1, Kallur Nava SARASWATHY2*, Khangembam Somibabu MEITEI2, Prakash Ranjan MONDAL2, Harpreet KAUR3, Kiran KUCHERIA1, Seema BHARDWAJ4, Gaznavi IDRIS1 1Department of Anatomy, All India Institute of Medical Sciences, Delhi 110029, India 2Department of Anthropology, University of Delhi, Delhi 110007, India 3INCLEN Trust International India, Delhi 110020, India 4University of Banswara, Banswara 327001, India Received 26 August 2010; accepted 11 January 2011 Abstract Rajasthan lies on the northwest border of India, and has acted as a major route for human movements since prehistoric and historic times. The present study was conducted to gain an insight into inter- and intrapopulation affinities or variations among the six population groups of Rajasthan. DNA samples from 221 unrelated individuals belonging to six endogamous population groups of Rajasthan, including both tribal (Bhil, Damaria, Garasia, Mina and Saharia) and caste populations (Rajput) were screened for 12 DNA (seven Alu indel and five RFLP) markers. All the loci were found to be polymorphic in all the studied populations. The GST values which determine the genetic differ- entiation between the population ranges from 1.5% in LPL to 8.7% in APO. This study suggests that the population groups of Rajasthan are genetically close to other Indian populations, reflecting a com- mon genetic unity among the Indian populations. The study also indicates European gene flow into the populations studied in this paper. Key words: Autosomal markers, common linguistic group, genetic affinity, genetic differentiation Introduction the Indus Valley civilization that is approximately 4500 years old, but disappeared about 3500 BP. The largest con- Rajasthan lies on the northwest border of India and has centration of Indus Valley civilization sites appear in an area varied topography ranging from Aravalli Hills (the oldest encompassing modern-day Rajasthan, Gujarat and Punjab. hill range in the world) to the Thar Desert, which covers In addition, the linguistic homogeneity of the region, includ- 61% of the land area of Rajasthan. The climate of Rajasthan ing both the caste and tribal groups of Rajasthan, is also ranges from scorching summers to chilly winters. This high- unique as they all fall within the Indo-European linguistic ly heterogeneous landscape could have geographically iso- family, which is mainly confined to the caste populations of lated the populations of Rajasthan and helped to maintain India. However, traces of the Dravidian linguistic group are their endogamous nature. Due to the harsh climatic condi- also found in the literature with respect to the Indus Valley tions, differential selective pressures are also likely to have civilization (Marshall, 1979; Knorozov et al., 1981). It is be- been present. The Indian subcontinent is considered as a lieved that speakers of Dravidian languages were wide- major corridor for the dispersal of modern humans as they spread throughout India before the arrival of Indo-European emerged out of Africa about 100000 years ago (Cavalli- speakers (Thapar, 1966). Subsequently, Indo-European Sforza et al., 1994; Cann, 2001). Since it is located at the speakers exercised dominance over the pre-existing pop- crossroads of Africa and West as well as East Eurasia ulations (Dravidian) and Dravidian speakers are currently (Cordaux et al., 2003), Rajasthan in India probably served as confined primarily to the southern part of India. These cul- one of the major land routes for human migrations. More- tural contacts and invasions have resulted in a high degree of over, from the fourth century BC onwards, India, particular- genetic and cultural differentiation among the people of ly in the north, was subjected for 2000 years to repeated Rajasthan. waves of penetration by alien peoples (Danielou, 2003; Indian populations are basically categorized into tribes Bhasin, 2006). The archaeological evidence for this is and castes. The tribes of India constitute only 8.2% of the present in the form of many prehistoric sites which include total Indian population and are more or less confined to rural India. The tribal populations of Rajasthan studied in this paper speak Indo-European languages as against the South * Correspondence to: Kallur Nava Saraswathy, Biochemical and Indian (Dravidian speakers) and East Indian (Austro-Asiatic Molecular Anthropological Laboratory, Department of Anthropol- ogy, University of Delhi, Delhi 110007, India. and Tibeto-Burman speakers) tribes of India. There are E-mail: [email protected] around 573 tribal populations in India, of which 75 are Published online 5 April 2011 categorized as primitive tribes by the Indian government. in J-STAGE (www.jstage.jst.go.jp) DOI: 10.1537/ase.100826 The caste group Rajput belongs to the major hierarchical © 2011 The Anthropological Society of Nippon 259 260 R. DADA ET AL. ANTHROPOLOGICAL SCIENCE varna system of India, comprising Brahmins, Kshatriyas, Materials and Methods Vaishyas, and Shudras. In this context the present study analyses five tribal populations (Bhils, Damaria, Garasia, Intravenous blood samples (5 ml) were collected from Minas, and Saharia) and one caste group of Rajasthan healthy, unrelated individuals of both sexes from six endog- (Rajputs) with respect to 12 autosomal markers. All the stud- amous groups (Bhils, Minas, Saharia, Garasia, Damaria, and ied populations belong to the Indo-European linguistic fam- Rajputs) of Rajasthan, with proper informed written consent. ily. Except for the Indo-European speaking Rajputs, which Samples were collected from the Banswara district of Rajas- is a caste group, all the other population groups are tribal. than (Figure 1) as this has the highest concentration of tribal Historically, the Bhils (Mann, 1987) and the Minas population (72.3%) (Census of India, 2001). This district (Bhasin, 2007) are taken to be the original inhabitants of shares borders with Gujarat and Madhya Pradesh states Rajasthan, constituting 39.3% and 50%, respectively, of the (Table 1). total tribal population of the state, whereas the Saharia is the only primitive tribe of Rajasthan, constituting 1% of the total Laboratory Analysis tribal population, and bears a resemblance to the Hindu caste populations of the area. Rajputs are thought to have originat- DNA was extracted following the salting-out protocol ed in Rajasthan only, but, according to James Tod (1832), (Miller et al., 1988). Seven human-specific Alu Indel (PV92, Rajputs have a Scythian origin—a stock which moved out of FXIIIB, D1, CD4, APO, ACE, mtNUC) and five restriction Caucasus towards the Indus Valley. The Garasias constitute fragment length polymorphism (RFLP) (NAT, LPL, PSCR, about 2.7% of the total tribal population and are thought to T2, ALB) markers were analyzed from the six population be descendants of Rajputs by Bhil women (Kapoor, 2002). samples. Amplification and analysis of DNA samples was The Damaria constitute 0.62% of the tribal population of the carried out using polymerase chain reaction (PCR), followed state and are taken to be a branch of the Bhils (Mehta, 2004). by restriction digestion as required. The primers and proto- In view of the diverse ethnic background of the popula- cols used for typing Alu Indel and mtNUC markers were as tions of Rajasthan vis-à-vis the numerous waves of migra- given by Stoneking et al. (1997), Majumder et al. (1999), tion in India, the main objective of the present paper is to and Watkins et al. (2001). The details of RFLPs studied are gain some insight into the peopling of India and to assess the given by the Allele Frequency Database (http://alfred.med. inter- and intra-population variation as well as the affinities yale.edu/alfred/index.asp). The PCR product for Alu Indel of the population groups of Rajasthan. markers and the digested PCR product for RFLP markers were run on a 2% agarose gel stained with EtBr (ethidium bromide) and then visualized in UV light for genotyping. Figure 1. Geographical location of the region (Banswara district) of sample collection. Vol. 119, 2011 GENOMIC STUDY AMONG RAJASTHAN POPULATIONS 261 Table 1. Anthropological information of the study populations Population Number Location of sampling Anthropological information Bhil 47 Banswara district Tribe, Indo-European (Bhili), numerically large, wide geographical distribution, among the earliest inhabitants of Rajasthan, trace their origin from Dravidian word for bow, ‘bilu,’ which is a characteristic weapon of Bhils Mina 31 Banswara district Largest tribal group of Rajasthan, Indo-European, among the earliest inhabitants of Rajasthan, claim to originate from Matsya Avatr. Saharia 30 Banswara district Tribe, Indo-European (Harauti), literal meaning—companion of tiger; clan names according to places of origin Garasia 53 Banswara district Tribe, thought to be descendents of Rajputs by Bhil women, Indo-European (dialect akin to Bhili) Damaria 26 Banswara district Tribe, Indo-European (Wagdi), taken to be a branch of the Bhils, migrated from Gujarat Rajputs 34 Banswara district Upper caste, Indo-European (Hindi), name derived from Sanskrit word ‘rajputra’ meaning son of a ruler Table 2. Allele frequencies for 12 autosomal markers in the six Statistical Analysis populations of Rajasthan Allele frequencies and heterozygosities at individual loci Populations as well as overall heterozygosity were calculated by using Marker the software POPGENE version 1.32 (Yeh and Yang, 1999). Bhil Rajput Saharia Mina Garasia Damaria Hardy–Weinberg equilibrium was tested using the chi Pv92 (+) 0.422 0.405 0.433 0.523 0.403 0.296 square (χ2) goodness of fit. Bonferroni’s correction for FXIIIB (+) 0.715 0.709 0.629 0.583 0.519 0.421 multiple comparisons was also applied. In order to evaluate D1 (+) 0.234 0.336 0.312 0.326 0.461 0.310 genetic differentiation among the studied groups, the locus- APO (+) 0.880 0.894 0.800 0.629 0.923 0.948 wise measure of genetic diversity (Nei, 1973) was calculated ACE (+) 0.510 0.613 0.578 0.416 0.730 0.544 using the software DISPAN (Ota, 1993).