Polymorphism of HLA Class I Genes in Meizhou Han Population Of

中国科技论文在线 http://www.paper.edu.cn

doi: 10.1111/j.1744-313X.2007.00669.x

PolymorphismBlackwell Publishing Ltd of HLA class I genes in Meizhou Han population of Guangdong, China

S. Chen,* W. Li,* Q. Hu,* Z. Liu,† Y. Xu* & A. Xu*

Summary Introduction Human leukocyte antigen (HLA) is an invaluable marker Han Chinese is the most popular ethnic group of China; for anthropological studies because of its extreme poly- Han inhabit all over the country. Guangdong (Canton) is morphism. Most of the studies carried out in Chinese located in southern China and Guangdong Han is a special populations are about HLA class II genes, but few about population in Han Chinese because of its particular class I genes. In the present study, we investigated HLA geographical location. People from northern China and class I polymorphism using polymerase chain reaction- the ethnic minorities of southern China both contributed sequencing-based typing (PCR-SBT) method in 104 to the formation of Guangdong Han, and thus Guangdong unrelated Han individuals in Meizhou of Guangdong, Han showed characteristics of various populations. The southern China. Twenty-three HLA-A, 43 HLA-B and genetic background of Guangdong Han is very useful 27 HLA-C alleles were identified and allele frequencies for understanding the formation of ethnic populations and two-locus (C/B) and three-locus (A/C/B) haplotypes in China. were statistically analysed. The most frequent HLA-A Meizhou is located in north-eastern Guangdong, allele is A*110101 with a frequency of 30.3%, followed and the major Han population in Meizhou is Hakka, by A*24020101 (22.2%) and A*2420 (11.6%). Among one of the three most predominant groups in Guangdong the 43 detected HLA-B alleles, B*5801 (17.0%), Han and even in the Han of southern China. The B*400101 (15.5%) and B*4601 (10.0%) were frequently Hakka distribute all over the world with a population observed. Among the 27 detected C alleles, the most more than 60 million. The formation of Hakka is a predominant one is Cw*07020101 (25.8%), followed debated issue, and for this reason the genetic study about by Cw*0717 (14.7%). The most frequent HLA-C/B Meizhou Han (Hakka) is especially important in the two-locus haplotype is Cw*07020101/B*400101 (10.1%). elucidation of the origin of Hakka and migration of The most common HLA-A/C/B three-locus haplotype Han Chinese and its association with other ethnic in Meizhou Han is A*110101/Cw*07020101/B*400101 groups. (3.4%). Phylogenetic tree based on HLA class I allele According to the historical evidences, the most popular frequencies genetically suggested that Meizhou Han has hypothesis about the origin of Meizhou Han is that the an affinity to southern Asian populations. The result may ancestors of the Meizhou Han originally emigrated from also reflect an admixture of Han and ethnic minorities the Central Plain to the south-eastern China in about 214 bc. of southern China. After that, they continuously immigrated to Guangdong. After five large migrations and admixtures with the local ethnic minorities, some of them finally settled down in Meizhou of Guangdong (Fang & Jiang, 1996) and became the Meizhou Han nowadays. Human leukocyte antigen (HLA) is a highly polymorphic * State Key Laboratory for Biocontrol, Guangdong Key Laboratory of genetic system in humans and plays an important role Pharmaceutical Functional Genes, Department of Biochemistry, in the immune response by presenting peptides to T College of Life Sciences, Sun Yat-sen (Zhongshan) University, Guangzhou, Guangdong, 510275 China and † Present address: lymphocytes (Parham, 1996). Because of the high degree Research Center for Molecular Biology, Jinan University, Guangzhou, of polymorphism, analysis of the HLA allele distribution Guangdong, 510630 China is valuable to trace the migration of ancient human Received 20 November 2006; revised 20 November 2006; accepted populations in anthropology studies (Imanish et al., 27 December 2006 1992) and many investigations have been done to Correspondence: Anlong Xu, Department of Biochemistry, College of elucidate the origin and genetic relationship of different Life Sciences, Sun Yat-sen (Zhongshan) University, Guangzhou, ethnic groups. In the present study, the allelic polymor- Guangdong, 510275 China. Tel: +86 20 8411 3655; phism of HLA-A, -B, and -C genes in Meizhou Han of Fax: +86 20 8403 8377; E-mail: [email protected] Guangdong was investigated by the method of polymerase These authors (S. Chen, W. Li) contributed equally to this work. chain reaction–sequencing-based typing (PCR-SBT). The

132 S. Chen et al.

43 HLA-B and 27 HLA-C alleles were detected, respectively (Table 1). In the HLA-A locus, A*110101 with a frequency of 30.3% is the most frequent allele in Meizhou Han. This allele was observed with a signiﬁcantly high frequency in Asians, such as Taiwanese (Shaw et al., 1999; Lin et al., 2000; Chu et al., 2001), Filipinos, Thais (26.0–40.0%), Malays (23.2%) (Bugawan et al., 1999), the Baloch tribe of Iran (14.2%) (Farjadian et al., 2004) and Northern Han in China (16.2%) (Hong et al., 2005). A*24020101 (22.2%) and A*2420 (11.6%) are another two frequent alleles in Meizhou Han. A*24020101 is also predominant in Asians. A*2402 was detected frequently in Japanese (37.9%) (Saito et al., 2000), Northern Han (15.2%) (Hong et al., 2005), Tibetans (27.2%) (Chen et al., 2006) and Taiwanese (16.6–86.3%) (Shaw et al., 1999; Lin et al., Figure 1. A map showing Meizhou Han and Taiwanese population 2000; Chu et al., 2001) in previous studies. A*2420 was location. (1) Meizhou Han; (2) populations from Taiwan. initially identiﬁed in Atayal of Taiwan (Scheltinga et al., 2000) and less frequent in northern population of China (Hong et al., 2005; Chen et al., 2006). The three common alleles accounted for 64.1% allele frequency in this locus. two-locus (C/B) and three-locus (A/C/B) haplotypes There are three alleles with a frequency of more than were analysed and the phylogenetic relationship of 10% in the HLA-B locus and B*5801 (17.0%) is the most the Meizhou Han with other populations was also frequent allele. Similar to what we observed in HLA-A discussed. locus, three frequent HLA-B alleles in Meizhou Han are also common in other Asian populations. B*5801 is a high frequency allele in some Chinese populations such as Materials and methods Hakka (10.2%) and Minnan (8.8%) of Taiwan (Shaw Peripheral blood samples were obtained from 104 unrelated, et al., 1999), Guangdong Han Chinese (8.1%) (Li et al., healthy individuals of Han population in Meizhou of 2006) and Singapore Chinese (10.4%) (Middleton et al., Guangdong, southern China (Fig. 1). PCR-SBT method 2004). B*4001 was frequently detected in Taiwanese was used for typing of the HLA-A, -B and -C (Pera et al., (21.3% to 34.9%) (Shaw et al., 1999), Guangdong Han 2000; Petersdorf & Malkki, 2000; Pozzi et al., 2000). The Chinese (14.4%) (Li et al., 2006) and Singapore Chinese HLA-A, -B and -C polymorphism was investigated in 99, (17.2%) as well (Middleton et al., 2004). B*4601, con- 100 and 99 individuals, respectively, because of failure of sidered as a unique allele in Asians, is the third frequent amplifying some loci for some samples. allele in Meizhou Han. This allele is predominant in Chinese Allele frequencies of HLA-A, -B and -C genes in Meizhou such as Taiwan Minnan (15.2%) (Shaw et al., 1999; Han were calculated by direct counting. The haplotype Lin et al., 2000), Singapore Chinese (14.5%) (Middleton frequencies were estimated using arlequin 2.0 (Schneider et al., 2004) and Guangdong Han Chinese (14.5%) (Li et al., 2000) with the Expectation-Maximization et al., 2006), but is rare in other populations of the world (EM) method. A dendrogram was constructed by the with a frequency of less than 0.1% in Europeans. In neighbour-joining method (Saitou & Nei, 1987) using the addition, HLA-B*1307 N, a rarely found allele, was phylip software (Felsenstein, 1993) based on the allele detected with a frequency of 8.5%. This probably frequencies of HLA-A, -B and -C genes in Meizhou Han, resulted from different typing approaches because the 11 populations in Taiwan (Ami, Atayal, Bunun, Hakka, previous data were rarely established from SBT. Although Minnan, Paiwan, Pazeh, Rukai, Thao, Toroko and Yami) 43 alleles were detected in this locus, the frequency of and 13 referenced populations out of China (Filipino, each allele out of 28 low frequency alleles is not more Korean, Malay, Thai, Australian, Arab Druze, Bulgarian, than 1%. Irish, Czech, Mexican, North American, Kenyan and In the HLA-C locus, Cw*07020101 (25.8%) and Zulu). Cw*0717 (14.7%) are the most frequent two alleles. According to previous studies, Cw*0702 distributed widely in worldwide populations including Chinese Results and discussion (20.8%), Thai (19.0%), Filipino (33.0%), Corsican (32.8%) (Grimaldi et al., 2002) and South American (32.8%) Allele frequencies (http://www.ncbi.nlm.nih.gov/projects/mhc/ihwg.cgi). In the present study, the diversity of HLA-A, -B, and -C Cw*07 is one of the most frequent alleles in Guang- genes in Meizhou Han of Guangdong was investigated dong Han (18.9%) (Bao et al., 2004; Ma et al., 2004). using high-resolution PCR-SBT method, and 23 HLA-A, There are seven HLA-C alleles with a frequency higher

Distribution of HLA class I alleles in Meizhou Han 133

Table 1. Allele frequencies of HLA-A, -B, and –C genes in Meizhou Han

Freqency (%) Frequency (%) Frequency (%) Allele (2n = 198) Allele (2n = 200) Allele (2n = 198)

A*110101 30.3 B*5801 17.0 Cw*07020101 25.8 A*24020101 22.2 B*400101 15.5 Cw*0717 14.7 A*2420 11.6 B*4601 10.0 Cw*030401 8.1 A*1112 8.6 B*1301 8.5 Cw*080101 8.1 A*1106 5.1 B*1307 N 8.5 Cw*030201 7.6 A*1102 4.6 B*4014 5.5 Cw*0102 6.6 A*3303 4.0 B*1502 3.5 Cw*0803 5.6 A*010101 1.5 B*15010101 3.0 Cw*0316 4.6 A*020101 1.5 B*4011 2.0 Cw*04010101 3.0 A*2423 1.5 B*520101 2.0 Cw*0309 2.0 A*0203 1.0 B*1508 1.5 Cw*0708 2.0 A*2428 1.0 B*4023 1.5 Cw*0410 1.5 A*2601 1.0 B*5401 1.5 Cw*0710 1.5 A*2616 1.0 B*5603 1.5 Cw*0403 1.0 A*3301 1.0 B*1302 1.5 Cw*0404 1.0 A*0103 0.5 B*1515 1.0 Cw*0602 1.0 A*0212 0.5 B*2704 1.0 Cw*070101 1.0 A*0213 0.5 B*3532 1.0 Cw*120201 1.0 A*0222 0.5 B*4036 1.0 Cw*0103 0.5 A*0227 0.5 B*4038 1.0 Cw*0407 0.5 A*0230 0.5 B*5110 1.0 Cw*0603 0.5 A*0238 0.5 B*510102 1.0 Cw*0606 0.5 A*020102 0.5 B*1505 0.5 Cw*0713 0.5 B*1522 0.5 Cw*0714 0.5 B*1524 0.5 Cw*1507 0.5 B*1539 0.5 Cw*120203 0.5 B*1553 0.5 Cw*150201 0.5 B*3511 0.5 B*3805 0.5 B*4002 0.5 B*4037 0.5 B*4803 0.5 B*5122 0.5 B*5301 0.5 B*5502 0.5 B*5507 0.5 B*5512 0.5 B*5602 0.5 B*5604 0.5 B*5806 0.5 B*510101 0.5 B*510202 0.5 B*520103 0.5

than 5% in Meizhou Han, accounting for 76.5% allele ihwg.cgi). The frequency in four populations including frequency. Yami (2%), Ami (4.1%), Rukai (6%) and Paiwan (6.9%) In general, the distribution of the frequent alleles of is similar with ours or lower than ours. On the other hand, HLA class I genes suggests that Meizhou Han has a close B*5801, the most common HLA-B allele in Meizhou relationship with the south-eastern Asians. Interestingly, Han, seems a low frequency allele in Chinese from northern we notice that the frequency of some alleles seems extremely China (Hong et al., 2005; Chen et al., 2006), but it is the low or high. This may represent the HLA characteristics third frequent allele in Chaoshan (10.69%) who are in this population and its complex genetic background. thought to be descended from the same ancestors as For example, the frequency of A*02 is 6%; this allele was Meizhou Han (Hu et al., 2007). Cw*07, also the most frequently detected in Chinese from northern China common HLA-C allele in Meizhou Han, varies from (Hong et al., 2005; Chen et al., 2006) and south-west 10.7% to 31.8% in 11 Taiwan populations and is the China (Shi et al., 2006). However, the frequency of A*02 most frequent allele in Atayal and the second frequent varies from 2% to 29.4% in 11 Taiwan populations we allele in Toroko (31.8%), Hakka (23.6%) and Minnan referenced (http://www.ncbi.nlm.nih.gov/projects/mhc/ (21.6%). The frequency of Cw*07 is as high as 77% in a

134 S. Chen et al.

relative isolated population, Drung (Chen et al., 2007). Table 2. HLA-C/B and HLA-A/C/B haplotypes in Meizhou Han Our results may imply that the Meizhou Han has a close (frequency > 0.01) relationship with Taiwanese and support the hypothesis that Taiwanese came from the coastal southern China. Haplotypes Frequency However, we can not exclude that the discrepancy with HLA-C/B the published data is partially due to different typing Cw*07020101/B*400101 0.101 approaches as we mentioned above that the most previous Cw*07020101/B*5801 0.048 HLA class I data in Chinese were not established from Cw*030401/B*1301 0.038 SBT. Cw*030201/B*5801 0.034 Cw*0717/B*4014 0.029 Cw*0717/B*5801 0.029 Haplotypes in Meizhou Han Cw*07020101/B*1307 N 0.024 Cw*030401/B*1307 N 0.019 HLA-C/B two-locus and HLA-A/C/B three-locus Cw*0102/B*4601 0.019 haplotypes (frequency > 1.0%) in Meizhou Han were Cw*080101/B*4601 0.019 listed in Table 2. The most frequent HLA-C/B two- Cw*0309/B*1307 N 0.014 locus haplotype is Cw*07020101/B*400101 (10.1%). Cw*0803/B*1502 0.014 Cw*0717/B*400101 0.014 The frequent ﬁve HLA-A/C/B three-locus haplotypes in Cw*030201/B*520101 0.014 Meizhou Han are A*110101/Cw*07020101/B*400101 Cw*0717/B*1301 0.010 (3.4%), A*2420/Cw*07020101/B*400101 (2.9%), Cw*04010101/B*15010101 0.010 A*110101/Cw*07020101/B*5801 (2.7%), A*3303/ Cw*0803/B*15010101 0.010 Cw*030201/B*5801 (2.4%) and A*110101/Cw*080101/ Cw*07020101/B*1502 0.010 B*4601 (1.9%). A*1101/Cw*0702/B*4001 haplotype Cw*080101/B*1502 0.010 was usually detected in Chinese such as Atayal (5.5%) Cw*07020101/B*3532 0.010 Cw*0102/B*4011 0.010 and Chinese Han (2.9%) and South Americans such Cw*0717/B*4038 0.010 as Brazilians (2.2%) and Guaranis (2.2% to 8.3%). Cw*0316/B*4601 0.010 A*3303/Cw*0302/B*5801 was frequently detected Cw*0410/B*4601 0.010 in Chinese Han (7.1%), Atayals (3.6%), Malays Cw*0102/B*5401 0.010 (5.0%), North Americans (5.4%) and South Americans Cw*0316/B*5801 0.010 (8.2%). Cw*04010101/B*5801 0.010 Cw*080101/B*5801 0.010

HLA-A/C/B Phylogenetic tree A*110101/Cw*07020101/B*400101 0.034 The phylogenetic tree based on the allele frequencies A*2420/Cw*07020101/B*400101 0.029 of the HLA-A, -B and -C of 25 populations clearly A*110101/Cw*07020101/B*5801 0.027 A*3303/Cw*030201/B*5801 0.024 indicated that Meizhou Han was clustered with 11 A*110101/Cw*080101/B*4601 0.019 Taiwan ethnic groups, suggesting their close genetic rela- A*110101/Cw*07020101/B*1301 0.014 tionship (Fig. 2). Among those 11 populations, Minnan A*110101/Cw*030401/B*1307 N 0.014 and Hakka are the two closest ethnic groups to Meizhou A*1112/Cw*0717/B*5801 0.014 Han. Eastern Asian has been classiﬁed into northern A*24020101/Cw*030401/B*1301 0.014 and southern populations (Ding et al., 2000). Distribution A*24020101/Cw*07020101/B*400101 0.014 A*020101/Cw*030401/B*1307 N 0.010 of HLA-A, -B and -C genes in Meizhou Han appears to A*110101/Cw*0410/B*15010101 0.010 be similar with that in southern population of East Asia, A*110101/Cw*0803/B*1502 0.010 suggesting that Meizhou Han belongs to the southern A*110101/Cw*07020101/B*3532 0.010 group of Chinese and may result from an admixture A*110101/Cw*0316/B*5801 0.010 of Han with ethnic minorities of southern China. In con- A*1102/Cw*04010101/B*4601 0.010 clusion, Meizhou Han probably has the same ancestor A*1112/Cw*0309/B*1301 0.010 A*24020101/Cw*0102/B*4011 0.010 as the other south-eastern Asian populations. It also A*24020101/Cw*0717/B*4014 0.010 supported the hypothesis that Taiwanese came from A*24020101/Cw*0102/B*4601 0.010 the coastal southern China and may be one of the A*24020101/Cw*0404/B*5801 0.010 ancestral sources of the Austronesian populations (Shaw A*2420/Cw*07020101/B*1307 N 0.010 et al., 1999; Lin et al., 2000; Su et al., 2000; Chu et al., A*2420/Cw*080101/B*5801 0.010 2001).

Acknowledgements The research was supported by Outstanding Young the key project (no. 021691) of Guangdong Natural Scientist Award (no. 39725007), the projects (no. 30178073 Foundation, and the project (no. 2001AA224021-04) and 30100275) and the science key project (no. 69935020) of the State High-Tech Development Project of the of the National Natural Science Foundation of China, Ministry of Science and Technology.

Distribution of HLA class I alleles in Meizhou Han 135

Figure 2. Neighbour-joining dendrogram showing the relationship between Meizhou Han and other 24 ethnic groups worldwide.

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