Blood genetic markers in Sri Lankan populationsЧreappraisal of the legend of Prince Vijaya.код для вставкиСкачать
AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 76:217-225 (1988) Blood Genetic Markers in Sri Lankan PopulationsReappraisal of the Legend of Prince Vijaya N. SAHA Department of Physiology, Faculty of Medicine, National University of Singapore, Singapore 0511 KEY WORDS Sri Lanka, Sinhalese, Tamils, Muslims, Serum proteins, Hemoglobin, Red cell enzymes, Electrophoresis, Isoelectric focusing, Population genetics ABSTRACT Serum protein (haptoglobin types; transferrin and group-specific component subtypes); haemoglobin and red cell enzymes (acid phosphatase, esterase D, glyoxalase I, 6-phosphogluconate dehydrogenase, adenylate kinase, and phosphoglucomutase (locus 1)(subtypes) were studied in the Sinhalese, Tamils, and Muslims of Sri Lanka. The allelic frequencies of all the polymorphic systems were similar in these populations without any significant differences. A close look at the present results and earlier investigations on 13 polymorphic loci controlled by 37 alleles did not reveal any genetic characteristics in the present-day Sinhalese population that are distinct from those in the. Tamils of Sri Lanka. As such, genetic evidence linking the legendary origin of the Sinhalese population to East India (Prince Vijaya) is lacking. INTRODUCTION The populations of present-day Sri Lanka are extremely diverse, with the Sinhalese constituting approximately 70% of the population. The remaining population includes the Tamils, Muslims, and other minorities, like the aboriginal Veddahs and the Burghers. Both myth and mystery surround the origin of the Sinhalese, who claim to be descendants of the peoples of North-East India more than 2,000 years ago, according to the legend of Prince Vijaya (Geiger, 1964).There have been immigrations from North-East India (present-day West Bengal and Bangladesh) to Sri Lanka for centuries. However, Basham (1964)holds that the Sinhalese came from the western coast of India. The origin of the Sinhalese language has been described as basically northwestern with considerable northeastern influence (Geiger, 1938) of the Indic languages and as quite distinct from the Dravidian languages. The Tamils originally came from Southern India (presently Tamilnadu) and migrated to Sri Lanka in the remote past, as evidenced by archeological discoveries in the northern part of the country, which was ruled by a 0 1988 ALAN R. LISS, INC series of monarchies. There have also been more recent Tamil immigrants from southern India, who came as workers during the British rule of the subcontinent and settled in Sri Lanka. Many of the Muslims of Sri Lanka claim Arab ancestry; they came as traders to the Malabar coast of India (present-day Kerala) (Fig. 1). A small group of Burghers are descendants of Dutch and Portuguese who intermarried with the local population. The main population groups, Sinhalese and Tamils, are quite distinct in their social habits, religion, and language-but intermarriage between the two groups has been going on for generations. Questions about the origin of the Sinhalese have interested many authors, because this Indo-European-speaking Buddhist population is thought to be as culturally (and presumably biologically) distinct from the Dravidian-speaking Hindu Tamils as from the minority remnants of the Reeeived January 8, 1981; revision accepted September 24, 1987. Address reprint requests to Dr. N. Saha, Department of Biochemistry, Faculty of Medicine, National University of Singapore, Singapore 0511. 218 N. SAHA I f Bombov \ R Fig. 1. Different regions of India and Sri Lanka associated with the historical background of Sri Lankan populations. aboriginal Veddahs of Sri Lanka. However, the Sinhalese system of kinship classification, cousin marriage, and the style of cooking, dress and housing are similar to those of Dravidian-speaking areas of southern India. Recent ethnographic work suggests that the cultural differences between the Sinhalese and Tamils and the Veddahs were overemphasized by earlier ethnographers (Brow, 1978). Genetic studies on the populations in Sri Lanka have been mostly limited to blood groups and abnormal hemoglobins in the past (De Silva, 1957; Wickremasinghe and Pon- nuswamy, 1963; Wickremasinghe et al., 1963). The first detailed study of genetic markers in the S r i Lankan populations was carried out by Kirk et al. , followed a decade later by a study on red cell enzymes in the Sinhalese population (Roberts et al., 1972). Kirk (1976) first tested the legendary origin of the Sinhalese by genetic distance analysis and concluded that the Sinhalese are nearer to the Bengalis and the Tamils than to the western Indians. Since the study by Kirk et al. (1962), no comparative study has been carried out among the Sinhalese and Tamil populations 'Abbreviations: obs. Groupspecific components Transferrin 1-1 2-1 2-1M 2-2 0 All Haptoglobin = observed; exp. = 22 IF2 IS2 2 2 All IF IFIS IS Y c2-1 c2-2 C3-1 C3-2 c3-3 C1-B C2-B C1-D C2-D c1-l 22 Phenotypes Systems expected. 25.00 14.00 11.00 27.00 18.00 8.00 0 103.00 7.80 1.00 30.00 1.00 72.00 0 104.00 1.42 71.00 27.00 5.00 1.00 0 0 1.00 0 1.00 0 106.00 1.93 20.10 23.85 7.08 26.94 15.99 9.03 0.00 102.99 69.77 30.01 3.23 0.81 0.17 0.00 0.81 0.18 0.81 0.17 105.96 104.01 - 73.62 2.62 27.77 Sinhalese No. No. obs. exp. Tamils 19.00 13.00 15.00 32.00 14.00 8.00 1.00 102.00 13.60 0 0 0 1.00 101.00 5.44 0 0 68.00 1.00 101.00 0.00 65.00 29.00 3.00 3.00 16.89 23.19 7.96 25.63 17.60 9.73 0.31 101.31 0.79 0.18 101.00 - 64.96 28.87 3.21 2.41 0.56 0.02 68.06 1.00 101.00 - ~ No. - exp. 3.06 3.00 28.88 29.00 No. obs. 13.00 13.00 7.00 5.00 7.00 2.00 0 47.00 2.75 10.30 15.92 6.15 7.50 5.79 1.36 0.00 47.02 0.76 0.21 46.97 - - 49.00 49.00 1.36 29.00 13.00 3.00 0 1.00 0 0 0 1.00 0 47.00 5.03 27.58 15.32 2.13 0.76 0.21 36.00 1.00 12.00 35.00 14.00 0 Muslims No. No. obs. exp. GC2 GC2 GC'F GC'S TfB TfD HP' HP2 HPO Allele TABLE 1. Distribution of serum protein polymorphism in Sri Lankan populations' - 0.4417 0.2621 0.2961 0.8113 0.1745 0.0047 0.0047 0.0047 - 0.1587 0.8413 - 0.4069 0.2794 0.3088 0.0049 0.0049 0.8020 0.1782 0.0149 0.1750 0.8250 0.0099 - - 0.4681 0.3617 0.1702 0.0106 0.7660 0.2128 0.0106 - 0.1429 0.8571 Gene frequencies Sinhalese Tamils Muslims 220 N. SAHA of Sri Lanka. Several new genetic marker polymorphisms have been described since the work of the above authors and that of Roberts et al. (1972) on which no data on the Sri Lankan populations are available. Isoelectric focusing now makes it possible to carry out a detailed study of some blood genetic markers for population genetics, but to date no study of isoelectric variations has been reported in the Sri Lankan populations. In view of the above developments and the controversies over the origin of the major Sinhalese population of Sri Lanka, we present here the results of a genetic marker study in three Sri Lankan populations, Sinhalese, Tamils, and Muslims. MATERIALS AND METHODS A total of 260 unrelated male Sri Lankans, 106 Sinhalese, 105 Tamil Hindus, and 49 Muslims, formed the study sample. They were attending the Singapore Anti-Tuberculosis Association (SATA) Clinic for medical examinations required for their employment as foreign workers. Samples of blood were collected in EDTA tubes. Plasma and packed cells were separated immediately by centrifugation at 2,000 rpm for 30 minutes. Both the plasma and the packed cells were stored frozen at -80°C in aliquot portions. Hemoglobin and red cell enzymes, except the ones mentioned below, were examined by horizontal starch-gel electrophoresis with standard techniques. Serum proteins (haptoglobin, transferrin albumin, and ceruloplasmin) were studied by polyacrylamide gel electrophoresis by the Ashton and Braden buffer system. Red cell acid phosphatase and esterase D were phenotyped with fluorogenic stains. Red cell glyoxalase I polymorphism was examined by the iodine staining method. Serum transferrin was subtyped by isoelectric focusing at pH 3.5-10 and pH 5-7 after desialation with neuraminidase. Group-specific component subtypes were identified by isoelectric focusing at pH 4-6.5 using the sulfosalisylic acid precipitation method. Red cell phosphoglucomutase polymorphism was studied by starch-gel electrophoresis and isoelectric focusing at pH 5-7. The gene frequencies of the different polymorphic loci were estimated by gene counting. The Hardy-Weinberg equilibrium was calculated by the chi-square (x2)test. RESULTS AND DISCUSSION Serum proteins The distribution of serum protein phenotypes and gene frequencies in the Sinhalese, Tamils, and Muslims of Sri Lanka is presented in Table 1. No significant difference in gene frequencies for Hp types, transferrin, and group-specific component subtypes was observed between these three populations. The frequency of Hp' was 0.16 in the Sinhalese, 0.17 in the Tamils, and 0.14 in the Muslims. Kirk et al. (1962) observed a similar frequency of Hp' (0.17) in the Sinhalese; however, the Tamils of Sri Lanka in their series had a lower frequency of Hpl (0.13) compared to the frequency (0.17)observed in the present study. An even lower frequency of Hp' has been reported in the Tamils of South India (Ananthakrishnan and Kirk, 1969; Ananthakrishnan, 1972). The frequency of Hp' in the Bengalis has been reported to be similar (Das et al., 1974; Mukherjee et al., 1974; Papiha et al., 1975; Saha, 1986). A solitary example of Hpo was observed among the Tamils of the present series. The incidence of Hpo has been reported to vary from 1%to 2% in Sri Lankan populations (Kirk et al., 1962). The lower incidence of Hpo in the present series could be due to the use of a sensitive polyacrylamide gel electrophoresis technique in the present investigation in place of the conventional starch-gel electrophoresis used in earlier investigations. The frequencies of Tf and Gc subtypes were also similar in these three groups. The frequency of Tfc' was found to be 0.81, 0.80, and 0.76, respectively, in the Sinhalese, Tamils, and Muslims of Sri Lanka. A low frequency of Tf c 3 ,and T f Dwas also present in these three groups. Only one case of Tp in the heterozygous form was detected in the Sinhalese. A similar low incidence of Tp and T P in the Sinhalese and Tamils was observed by earlier workers (Papiha and Wastell, 1974). A much higher frequency of Tf (0.87)was observed in the Bengali Muslims of Bangladesh (Saha, 1986), while a lower frequency was reported in the populations of West Bengal (Reddy et al., 1984). A similar frequency of T f ' has also been reported in the Tamils of Madras (0.77) and of Singapore by Kamboh and Kirk (1983) and Saha and Tan (1983). The frequency of Gc (subtype) alleles also did not exhibit any significant difference among the Sinhalese and Tamils of Sri Lanka. However, the Muslims had a significantly higher frequency of GcIS (0.36) than that observed in the Sinhalese (0.26) and Tamils (0.28). The Muslims also had a lower frequency of Gc2 (0.17) compared to that in the Sinhalese (0.30) and Tamils (0.31). The 89 50 30 10 90 2 18 36 56 58 3 1 62 1.37 12.25 27.37 40.99 56.04 2.92 0.04 59.00 28.55 7.90 0.55 37.00 0 15 26 41 57 1 1 59 28 9 0 37 1-1 2-1 2-2 A AC C All 1-1 2-1 2-2 All All All 1-1 2-1 2-2 'Abbreviations: obs. = observed;exp. = expected. Adenylate kinase 6-Phosphogluconate dehydrogenase Glyoxalase I Esterase D CB All B 4.28 33.44 65.28 0.00 103.00 52.96 41.07 7.96 101.99 8 26 69 0 103 53 41 8 102 A AB 58 0 53 5 0 6 28 55 105 0 104 1 Acid phosphatase 104.01 1.97 0.01 106.00 104 2 0 106 A AE E All Haemoglobin 53.11 4.78 0.11 58.00 57.10 4.80 0.10 62.00 2.16 17.68 36.16 56.00 4.49 31.01 53.50 0.00 89.00 46.94 36.11 6.94 89.99 104.00 1.00 0.00 105.00 Tamils No. obs. No. exp. Phenotypes Systems Sinhalese No. obs. No. exp. 26.00 0.00 0.00 26.00 26 0 26 0 15.02 0.97 0.02 16.01 1.00 8.00 16.00 25.00 2.57 16.61 26.81 0.75 46.74 36.67 15.66 1.62 54.00 53.00 1.00 0.00 54.00 15 1 0 16 0 10 15 25 3 16 27 1 47 37 15 2 54 53 1 0 54 Muslims No. obs. No. exp. A@ AK~ PGD~ PGD~ 0.1216 0.8484 0.9746 0.0254 0.1829 0.8171 0.2794 ESD~ GLO~ GL@ 0.7206 E~D' P" Pb 0.2039 0.7961 0.00 0.0094 HbE Pa 0.9906 0.0446 0.9569 0.9597 0.0403 0.1964 0.8036 0.2778 0.7222 0.2247 0.7753 0.00 0.0048 0.9952 0.0000 1.0000 0.9688 0.0312 0.2000 0.800 0.1759 0.8241 0.2340 0.7553 0.0106 0.0093 0.9907 Gene frequencies Sinhalese Tamils Muslims HbA Allele TABLE 2. Distribution of huemoplobin and red cell enzyme Dolvmorphisms in Sri Lankan uouulations' 222 N. SAHA GcW frequency observed in the Sinhalese and Tamils of Sri Lanka is much higher than that reported in a limited number of Indian populations (Walter et al., 1984). i I I I I dI I L d I ! .! ! r I 1 I 4 I d I 7 I , , I 5 2 n G P : Haxmoglobin and red cell enzymes The distribution of haemoglobin and five polymorphic red cell enzymes in the Sinhalese, Tamils, and Muslims of Sri Lanka is presented in Table 2. A low frequency of HbE has been observed in all three populations without any significant difference. A similarly low frequency of HbE has been reported in the Tamils of South India and in the Bengalis (Saha and Banerjee, 1973; Ghosh et al., 1981). The Veddahs in Sri Lanka have been reported to have a high frequency of HbE (Kirk et al., 1962; Wickremasinghe et al., 1963). Wickremasinghe and Ponnuswamy (1963)found no HbE in more than 1,000 samples of Sinhalese, while De Silva et al. (1959) had reported a high frequency in one locality of Sri Lanka. The latter might have been due to admixture with Veddahs in that area. An important feature to note is that the incidence of HbE is very low and is present in both the Sinhalese and the Tamils of Sri Lanka. This suggests that HbE has originated with the Veddahs rather than being introduced from South or East Asia, in which case the incidence should have been greater in the Sinhalese, contrary to Kirk's (1976) suggestion. The frequency of red cell acid phosphatase alleles was also very similar among the three populations of Sri Lanka. The frequency of p a was found to be 0.20,0.22,and 0.23 in the Sinhalese, Tamils, and Muslims, respectively. Roberts et al. (1972)reported a slightly higher frequency of pa (0.25)in their series of Sinhalese. No pc was detected in either the Sinhalese or the Tamils. One example of AP CB was encountered in the Muslims. A low frequency of p" has been reported in the Sinhalese and Tamils by previous workers (Ananthakrishnan, 1972;Roberts et al., 1972).A similar frequency of AP alleles has also been reported in the Bengalis (Das et al., 1970, 1974; Saha, 1986). The frequency of EsD' was also the same in the Sinhalese and Tamils (0.721,while that in the Muslims was 0.82. We have observed a similar frequency of EsD' in Bengali Muslims (0.73) by (Saha 19861, while Papiha et al. (1975) reported a EsD' of 0.77 in Bengali Muslims. The frequencies of GLO alleles were also found to be similar in the three groups, and the frequency of GLO' was 0.19 in the Sin- 223 GENETIC MARKERS IN SRI LANKANS TABLE 4. Gene frequencies in Sinhalese, Tamils, and Bangalis (13 loci and 37 alleles)1 Sinhalese Bengalis Tamils ABO P 4 r (0.1880) (0.1650) (0.6470) (0.1760) (0.1940) (0.6300) (0.1844; 0.2369) (0.2070; 0.2634) (0.608S; 0.4996) (0.5780) (0.4220) (0.6170) (0.3830) (0.6537; 0.5603) (0.3463; 0.4397) (0.6500) (0.08201 (0.0380) (0.0130) (0.2170) (0.6339) (0.0508) (0.0122) (0.0263) (0.2768) (0.6706; 0.6730) (0.0792; 0.1315) (0.0550; 0.0277) (0.0341) (0.1611; 0.1509) 0.1587 (0.1710) 0.8413 (0.8290) 0.1750 (0.114; 0.133) 0.8250 (0.886; 0.867) 0.1736 (0.175; 0.170; 0.185) 0.8264 (0.825; 0.830; 0.815) 0.8113 0.1745 0.0047 0.0047 (0.003) 0.0047 (0.003) 0.8020 0.1782 0.0149 0.0000 0.0049 (0.0006) 0.8710 0.1250 - 0.4417 0.2621 0.2964 0.0094 0.4069 0.2794 0.3088 0.0048 (0.004) (0.752) 0.2521 (0.248) 0.0194 (0.01) 0.2039 (0.253) 0.7961 (0.734) - (0.013) 0.7206 (0.763) 0.1829 0.0254 (0.016) 0.1216 (0.083); x2, 4.8 0.6968 (0.739) 0.2873 (0.261) 0.2247 (0.268) 0.7753 (0.730) (0.001) 0.7222 0.1964 0.0403 (0.022) 0.0446 (0.063) 0.6810 (0.736) 0.3095 (0.264) 0.016 0.0048 MN M N Rh R1 RZ Ro r' r HP HPl H D ~ Pa P" PC ESD' GLO' PGD~ AK2 PGM: PGe PGi@ PGi@ 0.0040 (0.015; 0.012) 0.2403 (0.203; 0.199) 0.7519 (0.790; 0.801) 0.0078 (0.007) 0.7326 (0.771) 0.2077 (0.2127) 0.0250 (0.040) 0.0385 (0.087) 0.7286 (0.722) 0.2571 (0.272) (0.003) 0.0048 (0.003) 'Gene frequencies in parentheses have been computed from the literature. halese and Tamils and 0.20 in the Muslims. A similar frequency of GLO' (0.21) had been reported in the Bengalis and Tamils (Saha, 1986; Ghosh, 1977). The frequency of PGD' was 0.04 in the Tamils and 0.03 in the Sinhalese and Muslims of the present series. A similar frequency of PGD' had been reported earlier in the Sinhalese, Tamils, and Bengalis (Roberts et al., 1972;Ananthakrishnan, 1972; Das et al., 1970, 1974; Papiha et al., 1975; Saha, 1987). The frequency of & was found to be 0.12 in the Sinhalese and 0.04 in the Tamils. Roberts et al. (1972)reported a frequency of 0.08 in their series of Sinhalese. A higher frequency of A3? (0.08) has also been reported in the Tamils of South India (Ananthakrishnan, 1972). The higher A P frequency ob- served in the Sinhalese in the present series should be interpreted with caution since the size of the sample was rather small. Table 3 shows the distribution of PGM subtypes in the Sinhalese, Tamils, and Muslims of Sri Lanka. The frequencies of the PGM subtype were also found to be similar in these three populations. The frequency of PGM1+ was slightly higher in the Sinhalese (0.55) and Muslims (0.58)than in the Tamils (0.53), while the frequency of other alleles was similar in the three groups. The frequency of PG@+ was higher in the Sinhalese (0.02) than in the Tamils (0.005) and Muslims. A low frequency of PG@+ has been reported in the Bengalis (Saha, 1986). One example of a new variant @I 6.5) has been observed in the Tamils and has also been reported in 224 N. SAHA other Indian and East Asian populations (Saha: unpublished observation). Monomorphic system Serum albumin, lactate and malate dehydrogenase, phosphohexose isomerase, phosphoglucomutase (locus 21, and superoxide dismutase were monomorphic. The absence of the lactate dehydrogenase variant is to be noted, as this variation has been reported in many populations of India, particularly in Bengal. In the present investigation, however, it may be due to the small size of the sample. In Table 4 the gene frequencies of 13 loci (37 alleles) in Sinhalese, Tamils, and Bengalis have been computed from our investigations and from the literature. It is clear that the frequencies of these alleles in the Sinhalese and Tamils of Sri Lanka are similar and that they are not significantly different from those in the Tamils of India or the Bengalis. The population genetics studies (present and earlier ones) so far carried out in the Sinhalese have not revealed any significant differences in the gene frequencies at 13 polymorphic loci from those of the Tamils in Sri Lanka. The allelic frequencies in the Sri Lankan populations, in general, are similar to those in other Indian populations. Furthermore, the results of five Gm allotypes and two Km allotypes show that the presentday Sinhalese are closer to the Sri Lankan and Indian Tamils than to the Bengalis (Matsumoto et al., in press). The present and earlier investigations produced no evidence whatsoever that the Sinhalese are genetically nearer to Eastern Indian populations than to the Tamils or to other South Indian populations. Even though there has been some legendary connection of the Sinhalese with East Indian or East Asian populations through trade or social links, there is no evidence to suggest that the present-day Sinhalese population is in any way genetically distinct from the Tamils of Sri Lanka. This finding might be due to free intermarriage between the two populations for several generations in the past in spite of the fact that they practice considerably different religion, language, and social systems. As far as we can see, the genetic evidence falls short of supporting the legend that the Sinhalese are descendants of Prince Vijaya. ACKNOWLEDGMENTS The author is grateful to Dr. N.C. Sengupta, Medical Director of the Singapore Anti-Tuberculosis Association (SATA) for permission to carry out this work. The technical assistance of the staff of SATA, Madame C.H. Ho and Jumiah bte Basair and the secretarial help of Miss W.F. Chan are gratefully recorded. The project was generously supported by grants from SATA, the Singapore Turf Club, the Shaw Foundation, and the Singapore National Heart Association. LITERATURE CITED Ananthakrishnan, R (1972) Further studies on the distribution of some serum protein and enzyme groups in South India. Hum. Genet. 15:172-176. Ananthakrishnan, R, and Kirk, RL (1969) The distribution of some serum protein and enzyme group systems in two endogamous groups of S. India. Indian J. Med. Res. 57:lOll-1017. Basham, AL (1964) Studies in Indian history and culture. XIV:Prince Vijaya and the Aryanization of Ceylon. Sambodhi Publications, Calcutta, pp. 162-177. 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