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1.  The Grandest Genetic Experiment Ever Performed on Man? – A Y-Chromosomal Perspective on Genetic Variation in India 
We have analysed Y-chromosomal data from Indian caste, Indian tribal and East Asian populations in order to investigate the impact of the caste system on male genetic variation. We find that variation within populations is lower in India than in East Asia, while variation between populations is overall higher. This observation can be explained by greater subdivision within the Indian population, leading to more genetic drift. However, the effect is most marked in the tribal populations, and the level of variation between caste populations is similar to the level between Chinese populations. The caste system has therefore had a detectable impact on Y-chromosomal variation, but this has been less strong than the influence of the tribal system, perhaps because of larger population sizes in the castes, more gene flow or a shorter period of time.
PMCID: PMC2987567  PMID: 21103011
Y chromosome; genetic variation; Indian caste system; endogamy; population substructure
2.  Genetic affinities among the lower castes and tribal groups of India: inference from Y chromosome and mitochondrial DNA 
BMC Genetics  2006;7:42.
India is a country with enormous social and cultural diversity due to its positioning on the crossroads of many historic and pre-historic human migrations. The hierarchical caste system in the Hindu society dominates the social structure of the Indian populations. The origin of the caste system in India is a matter of debate with many linguists and anthropologists suggesting that it began with the arrival of Indo-European speakers from Central Asia about 3500 years ago. Previous genetic studies based on Indian populations failed to achieve a consensus in this regard. We analysed the Y-chromosome and mitochondrial DNA of three tribal populations of southern India, compared the results with available data from the Indian subcontinent and tried to reconstruct the evolutionary history of Indian caste and tribal populations.
No significant difference was observed in the mitochondrial DNA between Indian tribal and caste populations, except for the presence of a higher frequency of west Eurasian-specific haplogroups in the higher castes, mostly in the north western part of India. On the other hand, the study of the Indian Y lineages revealed distinct distribution patterns among caste and tribal populations. The paternal lineages of Indian lower castes showed significantly closer affinity to the tribal populations than to the upper castes. The frequencies of deep-rooted Y haplogroups such as M89, M52, and M95 were higher in the lower castes and tribes, compared to the upper castes.
The present study suggests that the vast majority (>98%) of the Indian maternal gene pool, consisting of Indio-European and Dravidian speakers, is genetically more or less uniform. Invasions after the late Pleistocene settlement might have been mostly male-mediated. However, Y-SNP data provides compelling genetic evidence for a tribal origin of the lower caste populations in the subcontinent. Lower caste groups might have originated with the hierarchical divisions that arose within the tribal groups with the spread of Neolithic agriculturalists, much earlier than the arrival of Aryan speakers. The Indo-Europeans established themselves as upper castes among this already developed caste-like class structure within the tribes.
PMCID: PMC1569435  PMID: 16893451
3.  Presence of three different paternal lineages among North Indians: A study of 560 Y chromosomes 
Annals of human biology  2009;36(1):46-59.
The genetic structure, affinities, and diversity of the 1 billion Indians hold important keys to numerous unanswered questions regarding the evolution of human populations and the forces shaping contemporary patterns of genetic variation. Although there have been several recent studies of South Indian caste groups, North Indian caste groups, and South Indian Muslims using Y-chromosomal markers, overall, the Indian population has still not been well studied compared to other geographical populations. In particular, no genetic study has been conducted on Shias and Sunnis from North India.
This study aims to investigate genetic variation and the gene pool in North Indians.
Subjects and methods
A total of 32 Y-chromosomal markers in 560 North Indian males collected from three higher caste groups (Brahmins, Chaturvedis and Bhargavas) and two Muslims groups (Shia and Sunni) were genotyped.
Three distinct lineages were revealed based upon 13 haplogroups. The first was a Central Asian lineage harbouring haplogroups R1 and R2. The second lineage was of Middle-Eastern origin represented by haplogroups J2*, Shia-specific E1b1b1, and to some extent G* and L*. The third was the indigenous Indian Y-lineage represented by haplogroups H1*, F*, C* and O*. Haplogroup E1b1b1 was observed in Shias only.
The results revealed that a substantial part of today’s North Indian paternal gene pool was contributed by Central Asian lineages who are Indo-European speakers, suggesting that extant Indian caste groups are primarily the descendants of Indo-European migrants. The presence of haplogroup E in Shias, first reported in this study, suggests a genetic distinction between the two Indo Muslim sects. The findings of the present study provide insights into prehistoric and early historic patterns of migration into India and the evolution of Indian populations in recent history.
PMCID: PMC2755252  PMID: 19058044
Paternal lineages; Y-chromosomal markers; North Indians; migration
4.  Genetic variation in South Indian castes: evidence from Y-chromosome, mitochondrial, and autosomal polymorphisms 
BMC Genetics  2008;9:86.
Major population movements, social structure, and caste endogamy have influenced the genetic structure of Indian populations. An understanding of these influences is increasingly important as gene mapping and case-control studies are initiated in South Indian populations.
We report new data on 155 individuals from four Tamil caste populations of South India and perform comparative analyses with caste populations from the neighboring state of Andhra Pradesh. Genetic differentiation among Tamil castes is low (RST = 0.96% for 45 autosomal short tandem repeat (STR) markers), reflecting a largely common origin. Nonetheless, caste- and continent-specific patterns are evident. For 32 lineage-defining Y-chromosome SNPs, Tamil castes show higher affinity to Europeans than to eastern Asians, and genetic distance estimates to the Europeans are ordered by caste rank. For 32 lineage-defining mitochondrial SNPs and hypervariable sequence (HVS) 1, Tamil castes have higher affinity to eastern Asians than to Europeans. For 45 autosomal STRs, upper and middle rank castes show higher affinity to Europeans than do lower rank castes from either Tamil Nadu or Andhra Pradesh. Local between-caste variation (Tamil Nadu RST = 0.96%, Andhra Pradesh RST = 0.77%) exceeds the estimate of variation between these geographically separated groups (RST = 0.12%). Low, but statistically significant, correlations between caste rank distance and genetic distance are demonstrated for Tamil castes using Y-chromosome, mtDNA, and autosomal data.
Genetic data from Y-chromosome, mtDNA, and autosomal STRs are in accord with historical accounts of northwest to southeast population movements in India. The influence of ancient and historical population movements and caste social structure can be detected and replicated in South Indian caste populations from two different geographic regions.
PMCID: PMC2621241  PMID: 19077280
5.  Population Differentiation of Southern Indian Male Lineages Correlates with Agricultural Expansions Predating the Caste System 
PLoS ONE  2012;7(11):e50269.
Previous studies that pooled Indian populations from a wide variety of geographical locations, have obtained contradictory conclusions about the processes of the establishment of the Varna caste system and its genetic impact on the origins and demographic histories of Indian populations. To further investigate these questions we took advantage that both Y chromosome and caste designation are paternally inherited, and genotyped 1,680 Y chromosomes representing 12 tribal and 19 non-tribal (caste) endogamous populations from the predominantly Dravidian-speaking Tamil Nadu state in the southernmost part of India. Tribes and castes were both characterized by an overwhelming proportion of putatively Indian autochthonous Y-chromosomal haplogroups (H-M69, F-M89, R1a1-M17, L1-M27, R2-M124, and C5-M356; 81% combined) with a shared genetic heritage dating back to the late Pleistocene (10–30 Kya), suggesting that more recent Holocene migrations from western Eurasia contributed <20% of the male lineages. We found strong evidence for genetic structure, associated primarily with the current mode of subsistence. Coalescence analysis suggested that the social stratification was established 4–6 Kya and there was little admixture during the last 3 Kya, implying a minimal genetic impact of the Varna (caste) system from the historically-documented Brahmin migrations into the area. In contrast, the overall Y-chromosomal patterns, the time depth of population diversifications and the period of differentiation were best explained by the emergence of agricultural technology in South Asia. These results highlight the utility of detailed local genetic studies within India, without prior assumptions about the importance of Varna rank status for population grouping, to obtain new insights into the relative influences of past demographic events for the population structure of the whole of modern India.
PMCID: PMC3508930  PMID: 23209694
6.  Social Affiliation and the Demand for Health Services: Caste and Child Health in South India * 
Journal of development economics  2007;83(2):256-279.
This paper assesses the role of social affiliation, measured by caste, in shaping investments in child health. The special setting that we have chosen for the analysis – tea estates in the South Indian High Range – allows us to control nonparametrically for differences in income, access to health services, and patterns of morbidity across low caste and high caste households. In this controlled setting, low caste households spend more on their children's health than high caste households, reversing the pattern we would expect to find elsewhere in India. Moreover, health expenditures do not vary by gender within either caste group, in contrast once again with the male preference documented throughout the country. A simple explanation, based on differences in the returns to human capital across castes in the tea estates is proposed to explain these striking results.
PMCID: PMC2098704  PMID: 18046465
Health; Human Capital; Networks; Caste; Gender; Household decisions
7.  Clonal reproduction and genetic caste differences in a queen-polymorphic ant, Vollenhovia emeryi 
Biology Letters  2006;2(3):359-363.
Most social Hymenoptera are characterized by simple haploid sex determination and environment-based caste differentiation. This appears to be strikingly different in the queen-polymorphic ant Vollenhovia emeryi. Almost all long- and short-winged queens from a population in Central Japan were homozygous at three microsatellite loci, whereas workers were mostly heterozygous, suggesting either a complex system of genetic caste determination or, more likely, the production of female sexuals from unfertilized eggs by thelytokous parthenogenesis and of workers from fertilized eggs. Furthermore, male genotypes were not compatible with those of the queens and had exclusively the paternal allele found in the sterile, heterozygous workers, probably because males are produced from fertilized eggs after the exclusion of maternal nuclear DNA as recently reported for Wasmannia auropunctata. The genus Vollenhovia might provide an interesting model system to trace the evolution of unusual caste and sex determination systems.
PMCID: PMC1686177  PMID: 17148403
ant; caste; genetic; parthenogenesis; Vollenhovia
8.  Kinship Institutions and Sex Ratios in India 
Demography  2010;47(4):989-1012.
This article explores the relationship between kinship institutions and sex ratios in India at the turn of the twentieth century. Because kinship rules vary by caste, language, religion, and region, we construct sex ratios by these categories at the district level by using data from the 1901 Census of India for Punjab (North), Bengal (East), and Madras (South). We find that the male-to-female sex ratio varied positively with caste rank, fell as one moved from the North to the East and then to the South, was higher for Hindus than for Muslims, and was higher for northern Indo-Aryan speakers than for the southern Dravidian-speaking people. We argue that these systematic patterns in the data are consistent with variations in the institution of family, kinship, and inheritance.
PMCID: PMC3000033  PMID: 21308567
9.  Genetic diversity in India and the inference of Eurasian population expansion 
Genome Biology  2010;11(11):R113.
Genetic studies of populations from the Indian subcontinent are of great interest because of India's large population size, complex demographic history, and unique social structure. Despite recent large-scale efforts in discovering human genetic variation, India's vast reservoir of genetic diversity remains largely unexplored.
To analyze an unbiased sample of genetic diversity in India and to investigate human migration history in Eurasia, we resequenced one 100-kb ENCODE region in 92 samples collected from three castes and one tribal group from the state of Andhra Pradesh in south India. Analyses of the four Indian populations, along with eight HapMap populations (692 samples), showed that 30% of all SNPs in the south Indian populations are not seen in HapMap populations. Several Indian populations, such as the Yadava, Mala/Madiga, and Irula, have nucleotide diversity levels as high as those of HapMap African populations. Using unbiased allele-frequency spectra, we investigated the expansion of human populations into Eurasia. The divergence time estimates among the major population groups suggest that Eurasian populations in this study diverged from Africans during the same time frame (approximately 90 to 110 thousand years ago). The divergence among different Eurasian populations occurred more than 40,000 years after their divergence with Africans.
Our results show that Indian populations harbor large amounts of genetic variation that have not been surveyed adequately by public SNP discovery efforts. Our data also support a delayed expansion hypothesis in which an ancestral Eurasian founding population remained isolated long after the out-of-Africa diaspora, before expanding throughout Eurasia.
PMCID: PMC3156952  PMID: 21106085
10.  A Shared Y-chromosomal Heritage between Muslims and Hindus in India 
Human genetics  2006;120(4):543-551.
Arab forces conquered the Indus Delta region in 711 A.D. and, although a Muslim state was established there, their influence was barely felt in the rest of South Asia at that time. By the end of the tenth century, Central Asian Muslims moved into India from the northwest and expanded throughout the subcontinent. Muslim communities are now the largest minority religion in India, comprising more than 138 million people in a predominantly Hindu population of over one billion. It is unclear whether the Muslim expansion in India was a purely cultural phenomenon or had a genetic impact on the local population. To address this question from a male perspective, we typed eight microsatellite loci and 16 binary markers from the Y chromosome in 246 Muslims from Andhra Pradesh, and compared them to published data on 4,204 males from China, Central Asia, other parts of India, Sri Lanka, Pakistan, Iran, the Middle East, Turkey, Egypt and Morocco. We find that the Muslim populations in general are genetically closer to their non-Muslim geographical neighbors than to other Muslims in India, and that there is a highly significant correlation between genetics and geography (but not religion). Our findings indicate that, despite the documented practice of marriage between Muslim men and Hindu women, Islamization in India did not involve large-scale replacement of Hindu Y chromosomes. The Muslim expansion in India was predominantly a cultural change and was not accompanied by significant gene flow, as seen in other places, such as China and Central Asia.
PMCID: PMC2590854  PMID: 16951948
Y-chromosomal polymorphism; India; Muslim; Hindu
11.  Taxonomic studies on genus Tetramorium Mayr (Hymenoptera, Formicidae) with report of two new species and three new records including a tramp species from India with a revised key  
ZooKeys  2012;11-35.
Two new species of Tetramorium Mayr, namely Tetramorium shivalikense sp. n. and Tetramorium triangulatum sp. n. are described. Tetramorium triangulatum sp. n. belongs to the inglebyi-species group and is described based on worker, queen and male caste, while Tetramorium shivalikense sp. n. belongs to the ciliatum-species group and is described based on worker caste only. Three species viz., Tetramorium caldarium (Roger), Tetramorium tonganum Mayr and Tetramorium urbanii Bolton represent first records from India. The male caste is described for the first time in the case of Tetramorium tonganum. Among these, Tetramorium caldarium is a tramp species which extends its limit to India as well. A revised key to the Indian ants of this genus is also provided herewith.
PMCID: PMC3409682  PMID: 22855638
Tetramorium; Myrmicinae; new species; tramp species; India; key
12.  Worker caste determination in the army ant Eciton burchellii 
Biology Letters  2007;3(5):513-516.
Elaborate division of labour has contributed significantly to the ecological success of social insects. Division of labour is achieved either by behavioural task specialization or by morphological specialization of colony members. In physical caste systems, the diet and rearing environment of developing larvae is known to determine the phenotype of adult individuals, but recent studies have shown that genetic components also contribute to the determination of worker caste. One of the most extreme cases of worker caste differentiation occurs in the army ant genus Eciton, where queens mate with many males and colonies are therefore composed of numerous full-sister subfamilies. This high intracolonial genetic diversity, in combination with the extreme caste polymorphism, provides an excellent test system for studying the extent to which caste determination is genetically controlled. Here we show that genetic effects contribute significantly to worker caste fate in Eciton burchellii. We conclude that the combination of polyandry and genetic variation for caste determination may have facilitated the evolution of worker caste diversity in some lineages of social insects.
PMCID: PMC2391184  PMID: 17638672
division of labour; polyandry; multiple mating; worker polymorphism; social insects
13.  Gender variation in morphological patterns of lip prints among some north Indian populations 
Personal identification is an integral part of forensic investigations. For the same, DNA profiling and fingerprints are the most commonly used tools. But these evidences are not ubiquitous and may not necessarily be obtained from the crime scene. In such a scenario, other physical and trace evidences play a pivotal role and subsequently the branches employed are forensic osteology, odontology, biometrics, etc. A relatively recent field in the branch of forensic odontology is cheiloscopy or the study of lip prints. A comparison of lip prints from the crime scene and those obtained from the suspects may be useful in the identification or narrowing down the investigation.
The purpose of the present study is to determine the gender and population variability in the morphological patterns of lip prints among brahmins, Jats, and scheduled castes of Delhi and Haryana, India.
Settings and Design:
Samples were collected from Jats, brahmins, and scheduled castes of Delhi and Haryana. The total sample size consisted of 1399 individuals including 781 males and 618 females in the age group of 8–60 years. Care was taken not to collect samples from genetically related individuals. The technique was standardized by recording lip prints of 20 persons and analyzing them.
Materials and Methods:
Lip prints were collected by using a corporate's invisible tape and analyzed using a hand lens. The patterns were studied along the entire length and breadth of both the upper and the lower lip. The data were analyzed by SPSS statistical package version 17 to determine the frequencies and percentages of occurrence of the pattern types in each population group and a comparison between males and females among the groups was carried out by using the z test.
Results and Conclusions:
The z-test comparison between patterns of males and females shows significant differences with respect to pattern types I’, II, III, and IV among brahmins; I’, II, III, IV, and Y among Jats; and I, I’, II, III, and V among scheduled castes. Thus, it can be concluded that the variability of the lip print pattern can help sex differentiation among groups and that more studies on the lip print pattern should be carried out to bring new dimensions to forensic anthropology and to aid the law enforcement agencies.
PMCID: PMC3470412  PMID: 23087577
Cheiloscopy; gender; population variability
14.  Influence of language and ancestry on genetic structure of contiguous populations: A microsatellite based study on populations of Orissa 
BMC Genetics  2005;6:4.
We have examined genetic diversity at fifteen autosomal microsatellite loci in seven predominant populations of Orissa to decipher whether populations inhabiting the same geographic region can be differentiated on the basis of language or ancestry. The studied populations have diverse historical accounts of their origin, belong to two major ethnic groups and different linguistic families. Caucasoid caste populations are speakers of Indo-European language and comprise Brahmins, Khandayat, Karan and Gope, while the three Australoid tribal populations include two Austric speakers: Juang and Saora and a Dravidian speaking population, Paroja. These divergent groups provide a varied substratum for understanding variation of genetic patterns in a geographical area resulting from differential admixture between migrants groups and aboriginals, and the influence of this admixture on population stratification.
The allele distribution pattern showed uniformity in the studied groups with approximately 81% genetic variability within populations. The coefficient of gene differentiation was found to be significantly higher in tribes (0.014) than caste groups (0.004). Genetic variance between the groups was 0.34% in both ethnic and linguistic clusters and statistically significant only in the ethnic apportionment. Although the populations were genetically close (FST = 0.010), the contemporary caste and tribal groups formed distinct clusters in both Principal-Component plot and Neighbor-Joining tree. In the phylogenetic tree, the Orissa Brahmins showed close affinity to populations of North India, while Khandayat and Gope clustered with the tribal groups, suggesting a possibility of their origin from indigenous people.
The extent of genetic differentiation in the contemporary caste and tribal groups of Orissa is highly significant and constitutes two distinct genetic clusters. Based on our observations, we suggest that since genetic distances and coefficient of gene differentiation were fairly small, the studied populations are indeed genetically similar and that the genetic structure of populations in a geographical region is primarily influenced by their ancestry and not by socio-cultural hierarchy or language. The scenario of genetic structure, however, might be different for other regions of the subcontinent where populations have more similar ethnic and linguistic backgrounds and there might be variations in the patterns of genomic and socio-cultural affinities in different geographical regions.
PMCID: PMC549189  PMID: 15694006
15.  Eight-year experience with 3392 endoscopically proven duodenal ulcers in Durban, 1972-79. Rise and fall of duodenal ulcers and a theory of changing dietary and social factors. 
Gut  1981;22(4):327-331.
From 1972 to 1979, a total of 3392 patients with endoscopically proven ulcers were seen, in six ethnic groups. The distribution was as follows: Africans 456 males, 182 females; Muslim Gujerati Indians 206 males, 60 females; Hindu Hindi 433 males; total North Indians 639 males, 195 females; Hindu Tamils 593 males, 184 females; Hindu Telegu 179 males, 46 females; total South Indians 872 males, 230 females, and Whites 465 males, 303 females. In the continent of India, it is predominantly the South Indians who suffer from duodenal ulcer. In Durban, the number of North Indians with duodenal ulcers approximates that of those from the South (North: South ratio 0.83). The first question raised by this study is that the protective factors in North Indians in India are not genetic, and are lost when they emigrate to Natal. This may be due to changes in diet. A seasonal analysis indicates that, for females, there is a striking Autumn and Winter predominance in all Indian groups, reaching 80% in Muslims and Telegus but not in African females (52.7%). The second question raised by this study is that protective factors must be sought which operate in Indian females in the Spring and Summer months. The third question emanating from this study is that duodenal ulcers (and ischaemic heart disease) appear to increase in times of dietary and social change. This occurred in the West from 1890 to 1960, and is still occurring in the Third World. The restoration of dietary fibre and unsaturated fat, and the possible adjustment to stress in the West since 1960, has been accompanied by a fall in the incidence of these diseases. A 'changing factors' theory of duodenal ulcers and ischaemic heart disease is proposed. These conditions fall when a 'plateau situation' is reached.
PMCID: PMC1419150  PMID: 7239324
16.  Canine tooth dimorphism: An adjunct for establishing sex identity 
Teeth are an excellent material for genetic, odontological and forensic investigations and research purpose. From all the teeth, the mandibular canines are found to exhibit sexual dimorphism. However, very few studies have been published on maxillary canine's measurements.
1. To find out utility of maxillary and mandibular canine width as a tool for sex determination in Central Indian population. 2. To find out the average size of canines in males and females of Central Indian population. 3. To compare the findings with National and International studies
Materials and Methods:
The present study was conducted in 100 cases in the age group of 17-21 years. Mesiodistal width of right and left mandibular and maxillary canines were measured on the casts with digital calliper and subjected to statistical analysis.
Statistical Analysis:
Statistical analysis was done to assess sex difference using Students “t” test (paired).
Results and Conclusions:
It was seen that a definite statistically significant sexual dimorphism exists when mandibular and maxillary canine measurements were compared. Thus, it can be suggested that canine width measurements can be used as an adjunct for sex identification purpose in Central Indian Population.
PMCID: PMC3669482  PMID: 23741147
Canine dimorphism; forensic odontology; sex determination
17.  Genetic polymorphism in leaf-cutting ants is phenotypically plastic 
Advanced societies owe their success to an efficient division of labour that, in some social insects, is based on specialized worker phenotypes. The system of caste determination in such species is therefore critical. Here, we examine in a leaf-cutting ant (Acromyrmex echinatior) how a recently discovered genetic influence on caste determination interacts with the social environment. By removing most of one phenotype (large workers; LW) from test colonies, we increased the stimulus for larvae to develop into this caste, while for control colonies we removed a representative sample of all workers so that the stimulus was unchanged. We established the relative tendencies of genotypes to develop into LW by genotyping workers before and after the manipulation. In the control colonies, genotypes were similarly represented in the large worker caste before and after worker removal. In the test colonies, however, this relationship was significantly weaker, demonstrating that the change in environmental stimuli had altered the caste propensity of at least some genotypes. The results indicate that the genetic influence on worker caste determination acts via genotypes differing in their response thresholds to environmental cues and can be conceptualized as a set of overlapping reaction norms. A plastic genetic influence on division of labour has thus evolved convergently in two distantly related polyandrous taxa, the leaf-cutting ants and the honeybees, suggesting that it may be a common, potentially adaptive, property of complex, genetically diverse societies.
PMCID: PMC2169285  PMID: 17301017
phenotype; reaction norm; Acromyrmex; caste; polyandry; genotype
18.  Fine-scale population structure and the era of next-generation sequencing 
Human Molecular Genetics  2010;19(R2):R221-R226.
Fine-scale population structure characterizes most continents and is especially pronounced in non-cosmopolitan populations. Roughly half of the world's population remains non-cosmopolitan and even populations within cities often assort along ethnic and linguistic categories. Barriers to random mating can be ecologically extreme, such as the Sahara Desert, or cultural, such as the Indian caste system. In either case, subpopulations accumulate genetic differences if the barrier is maintained over multiple generations. Genome-wide polymorphism data, initially with only a few hundred autosomal microsatellites, have clearly established differences in allele frequency not only among continental regions, but also within continents and within countries. We review recent evidence from the analysis of genome-wide polymorphism data for genetic boundaries delineating human population structure and the main demographic and genomic processes shaping variation, and discuss the implications of population structure for the distribution and discovery of disease-causing genetic variants, in the light of the imminent availability of sequencing data for a multitude of diverse human genomes.
PMCID: PMC2953744  PMID: 20876616
19.  Control of reproductive dominance by the thelytoky gene in honeybees 
Biology Letters  2007;3(3):292-295.
Differentiation into castes and reproductive division of labour are a characteristics of eusocial insects. Caste determination occurs at an early stage of larval development in social bees and is achieved via differential nutrition irrespective of the genotype. Workers are usually subordinate to the queen and altruistically refrain from reproduction. Workers of the Cape honeybee (Apis mellifera capensis) do not necessarily refrain from reproduction. They have the unique ability to produce female offspring parthenogenetically (thelytoky) and can develop into ‘pseudoqueens’. Although these are morphologically workers, they develop a queen-like phenotype with respect to physiology and behaviour. Thelytoky is determined by a single gene (th) and we show that this gene also influences other traits related to the queen phenotype, including egg production and queen pheromone synthesis. Using 566 microsatellite markers, we mapped this gene to chromosome 13 and identified a candidate locus thelytoky, similar to grainy head (a transcription factor), which has been shown to be highly expressed in queens of eusocial insects. We therefore suggest that this gene is not only important for determining the pseudoqueen phenotype in A. m. capensis workers, but is also of general importance in regulating the gene cascades controlling reproduction and sterility in female social bees.
PMCID: PMC2464700  PMID: 17412668
thelytoky; pleiotropy; Apis mellifera
20.  Developmental Pathways of Psammotermes hybostoma (Isoptera: Rhinotermitidae): Old Pseudergates Make up a New Sterile Caste 
PLoS ONE  2012;7(9):e44527.
Ergonomic efficiency is in termites maximized by task partitioning among specialized castes. The isopteran caste systems can be classified as either (i) linear, when tasks are performed by pluripotent immatures (pseudergates), retaining the ability to develop into winged imagoes or (ii) bifurcated, with the presence of a true worker caste, which diverges early and permanently from the sexual (nymph/alate) line.
Principal Findings
Here, we report on the ontogenetic potentialities of the highly polymorphic sand termite Psammotermes hybostoma. Beside numerous pluripotent pseudergates, constituting the main work force, some larger non-feeding apterous immatures, also occur. These individuals are unable to proceed to the winged imago stage, but store large amounts of fat and also give rise to large soldiers. Soldiers therefore originate from a wide range of apterous instars, consequently being highly polymorphic.
The caste system of P. hybostoma is essentially linear, as in other basal Rhinotermitidae, but is distinguished by the late bifurcation leading to large apterous immatures. Because these large worker-like individuals deviate late and do not perform worker tasks, they cannot be considered homologous to the true workers of Termitidae and advanced Rhinotermitidae, but they provide a novel example of the evolution of sterile immatures in termites.
PMCID: PMC3444481  PMID: 23028554
21.  Reconstructing Indian Population History 
Nature  2009;461(7263):489-494.
India has been underrepresented in genome-wide surveys of human variation. We analyze 25 diverse groups to provide strong evidence for two ancient populations, genetically divergent, that are ancestral to most Indians today. One, the “Ancestral North Indians” (ANI), is genetically close to Middle Easterners, Central Asians, and Europeans, while the other, the “Ancestral South Indians” (ASI), is as distinct from ANI and East Asians as they are from each other. By introducing methods that can estimate ancestry without accurate ancestral populations, we show that ANI ancestry ranges from 39-71% in India, and is higher in traditionally upper caste and Indo-European speakers. Groups with only ASI ancestry may no longer exist in mainland India. However, the Andamanese are an ASI-related group without ANI ancestry, showing that the peopling of the islands must have occurred before ANI-ASI gene flow on the mainland. Allele frequency differences between groups in India are larger than in Europe, reflecting strong founder effects whose signatures have been maintained for thousands of years due to endogamy. We therefore predict that there will be an excess of recessive diseases in India, different in each group, which should be possible to screen and map genetically.
PMCID: PMC2842210  PMID: 19779445
22.  Distribution of CC-chemokine receptor-5-∆32 allele among the tribal and caste population of Vidarbha region of Maharashtra state 
Genetic relationships among the ethnic groups are not uniform across the geographical region. Considering this assumption, we analyzed the frequency of the CC-chemokine receptor-5 (CCR5)-∆32 allele of the CCR5 chemokine receptor, which is considered a Caucasian marker, in Bhil tribal and Brahmin caste sample sets from the population.
108 blood samples were collected from 6 tribe's populations and a caste population from the district of Vidarbha region.
The presence of low frequencies of CCR5-Δ32 in an individual of Bhil tribe (0.034, χ2 value 0.017) in the present study implies that these communities may have a better resistance toward human immunodeficiency virus (HIV)/acquired immunodeficiency syndrome (AIDS) than the other studied tribe sample, as non-show such mutation.
The marginal presence of the allele seen in the studied tribal population could be due to gene flow from the people of European descent. However, lack of the homozygous CCR5-Δ32 mutation and the low prevalence of heterozygous CCR5-Δ32 mutations suggest that the Indians are highly susceptible to HIV/AIDS, and this correlates with the highest number of HIV/AIDS infected individuals in India.
PMCID: PMC3722632  PMID: 23901195
Allele frequency; CC-chemokine receptor-5-∆32; India; genetic polymorphism; tribes; Vidarbha
23.  Genomic view on the peopling of India 
India is known for its vast human diversity, consisting of more than four and a half thousand anthropologically well-defined populations. Each population differs in terms of language, culture, physical features and, most importantly, genetic architecture. The size of populations varies from a few hundred to millions. Based on the social structure, Indians are classified into various caste, tribe and religious groups. These social classifications are very rigid and have remained undisturbed by emerging urbanisation and cultural changes. The variable social customs, strict endogamy marriage practices, long-term isolation and evolutionary forces have added immensely to the diversification of the Indian populations. These factors have also led to these populations acquiring a set of Indian-specific genetic variations responsible for various diseases in India. Interestingly, most of these variations are absent outside the Indian subcontinent. Thus, this review is focused on the peopling of India, the caste system, marriage practice and the resulting health and forensic implications.
PMCID: PMC3514343  PMID: 23020857
Admixture; caste; Indians; mtDNA; tribe; Y-chromosome
24.  Environmental and genetic controls of soldier caste in a parasitic social wasp 
Scientific Reports  2012;2:729.
A larval army caste is found in some parasitic wasps with polyembryonic or clonal proliferation, where many clone larvae emerge from a single egg. In contrast to non-parasitic eusocial Hymenoptera, sterile soldier larvae that protect their clonal reproductives are found in both females and males. Recently, the proportion of soldier larvae has been found to vary radically, depending on the internal conditions of the host, such as multiparasitism by other larval parasites. However, the proportion of male soldier larvae is constant, irrespective of the host internal environment. It is unknown if these traits are heritable. Here we show that a high heritability is found in both sexes, while, in the 6th instar hosts, substantially lower heritability is found in females. These results imply that the structure of the larval caste is determined genetically by both female and male embryonic cells, but more likely modified environmentally in females.
PMCID: PMC3476458  PMID: 23087811
25.  Socioeconomic Inequalities in Childhood Undernutrition in India: Analyzing Trends between 1992 and 2005 
PLoS ONE  2010;5(6):e11392.
India experienced a rapid economic boom between 1991 and 2007. However, this economic growth has not translated into improved nutritional status among young Indian children. Additionally, no study has assessed the trends in social disparities in childhood undernutrition in the Indian context. We examined the trends in social disparities in underweight and stunting among Indian children aged less than three years using nationally representative data.
We analyzed data from the three cross-sectional rounds of National Family Health Survey of India from 1992, 1998 and 2005. The social factors of interest were: household wealth, maternal education, caste, and urban residence. Using multilevel modeling to account for the nested structure and clustering of data, we fit multivariable logistic regression models to quantify the association between the social factors and the binary outcome variables. The final models additionally included age, gender, birth order of child, religion, and age of mother. We analyzed the trend by testing for interaction of the social factor and survey year in a dataset pooled from all three surveys.
While the overall prevalence rates of undernutrition among Indian children less than three decreased over the 1992–2005 period, social disparities in undernutrition over these 14 years either widened or stayed the same. The absolute rates of undernutrition decreased for everyone regardless of their social status. The disparities by household wealth were greater than the disparities by maternal education. There were no disparities in undernutrition by caste, gender or rural residence.
There was a steady decrease in the rates of stunting in the 1992–2005 period, while the decline in underweight was greater between 1992 and 1998 than between 1998 and 2005. Social disparities in childhood undernutrition in India either widened or stayed the same during a time of major economic growth. While the advantages of economic growth might be reaching everyone, children from better-off households, with better educated mothers appear to have benefited to a greater extent than less privileged children. The high rates of undernutrition (even among the socially advantaged groups) and the persistent social disparities need to be addressed in an urgent and comprehensive manner.
PMCID: PMC2894973  PMID: 20617192

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