Distributions of HLA-A and -B alleles and haplotypes in the Yi ethnic minority of Yunnan, China: relationship to other populations

Distributions of HLA-A and -B alleles and haplotypes in the Yi ethnic minority of Yunnan, China: relationship to other populations 1 2 first-author 1 3 corresp ‡Corresponding Authors ‡Corresponding Authors †E-mail:gyang@mcg.eduwzy218@mail.xjtu.edu.cn †E-mail:gyang@mcg.eduwzy218@mail.xjtu.edu.cn gyang@mcg.edu 4 5 6 7 1 2 8 2 3 1 2 1 2 1 1 1 2 corresp ‡Corresponding Authors ‡Corresponding Authors †E-mail:gyang@mcg.eduwzy218@mail.xjtu.edu.cn †E-mail:gyang@mcg.eduwzy218@mail.xjtu.edu.cn gyang@mcg.edu 3 Department of Forensic Medicine, School of Medicine, Xi’an Jiaotong University, Xi’an 710061, China Department of Forensic Medicine, School of Medicine, Xi’an Jiaotong University, Xi’an 710061, China Key Laboratory of Environment and Genes Related to Diseases of Ministry of Education, Xi’an Jiaotong University, Xi’an 710061, China Key Laboratory of Environment and Genes Related to Diseases of Ministry of Education, Xi’an Jiaotong University, Xi’an 710061, China Vascular Biology Center, Medical College of Georgia, Augusta, Georgia 30912, USA Vascular Biology Center, Medical College of Georgia, Augusta, Georgia 30912, USA Shaan′xi Blood Center, Xi’an 710061, China Shaan′xi Blood Center, Xi’an 710061, China Institute of Molecular Medicine and Genetics, Medical College of Georgia, Augusta, Georgia 30912, USA Institute of Molecular Medicine and Genetics, Medical College of Georgia, Augusta, Georgia 30912, USA Department of Public Health, School of Medicine, Xi′an Jiaotong University, Xi′an 710061, China Department of Public Health, School of Medicine, Xi′an Jiaotong University, Xi′an 710061, China Center of Forensic Sciences, Beijing Genomics Institute, Beijing 101300, China Center of Forensic Sciences, Beijing Genomics Institute, Beijing 101300, China Faculty of Basic Medical Science, Nanchang University, Nanchang 330006, China Faculty of Basic Medical Science, Nanchang University, Nanchang 330006, China Objective: 2 To investigate the distributions of human leukocyte antigen (HLA)-A and -B alleles and HLA-A-B haplotypes in the Yi ethnic minority of the Yunnan Province, situated in southwestern China. Methods: 2 DNA typing for HLA-A and -B loci was performed using the polymerase chain reaction-sequence-based typing (PCR-SBT) method on 114 randomly selected healthy individuals of the Yi population. The allelic frequencies of HLA-A and -B loci were calculated by direct counting and HLA-A-B haplotypes were estimated using the expectation maximization algorithm. Results: 2 A total of 17 HLA-A and 38 HLA-B alleles were found in the Yi population. The most frequent alleles were A*2402 (32.46%), A*1101 (26.32%), and A*0203 (10.09%) at the HLA-A locus and B*4601 (12.28%), B*1525 (10.09%), B*4001 (8.77%), and B*3802 (7.89%) at the HLA-B locus. The predominant HLA-A-B haplotypes were A*2402-B*1525 (7.86%) and A*0203-B*3802 (5.64%), followed by A*1101-B*4001 (4.69%). Phylogenetic analysis indicates that the Yi population in the Honghe, Yunnan Province of China basically belongs to groups of southeastern Asian origin, but shares some characteristics with northeastern Asian groups. Conclusion: 2 The present study may add to the understanding of HLA polymorphism in the Yi ethnic group that was poorly defined previously, and provide useful information for bone marrow transplantation, anthropological research, and forensic sciences as well as for disease-association studies. Yi ethnic minority Human leukocyte antigen (HLA)-A HLA-B Allele Haplotype Polymerase chain reaction-sequence-based typing (PCR-SBT) Introduction 1 The major histocompatibility complex (MHC) is a genetic region defined initially by the rejection of skin grafts in genetically incompatible mouse strains. In humans, the human leukocyte antigen (HLA) region, equivalent to the MHC, comprises about 3 Mb located on the short arm of chromosome 6 (6p21.3) (Erlich et al., 2001 2001 ). HLA genes are divided into three basic groups: class I, class II, and class III. HLA-A, -B, and -C are the major genes in MHC class I, the most polymorphic and variable one, with currently more than 2 678 alleles identified (situation on July 2009) ( http://www.ebi.ac.uk/imgt/hla/stats.html http://www.ebi.ac.uk/imgt/hla/stats.html ). It has been shown that the distributions of HLA alleles and haplotypes are different from one ethnic group to another or even between the members of the same ethnic group living in different geographical areas (Shen et al., 2008 2008 ). Due to its high degree of polymorphisms and variability in allele/haplotype frequencies, HLA loci analyses have become a valuable tool to trace population migration and origin, and to study population relationships for anthropological purposes (Hu et al., 2007 2007 ). The Yi ethnic minority, previously named “Luoluo,” is a modern ethnic group seen in China, Vietnam, and Thailand. With 8 million people, the Yi ethnic minority represents the 7th largest of 55 ethnic minority groups officially recognized in China. There are 3300 Yi people (1999 census) living in northeastern Vietnam ( http://www.chinakindnesstour.com/Chinainfo/minority/EthnicMinority/Chinainfo128.shtml http://www.chinakindnesstour.com/Chinainfo/minority/EthnicMinority/Chinainfo128.shtml ). Nearly all the Yi people live in the mountainous areas of Sichuan, Yunnan, Guizhou, and Guangxi Provinces in southwestern China, usually inhabiting the sides of steep mountain slopes far from cities. In the period between the 2nd century B.C. and the early Christian era, the activities of the ancient Yi centered on the areas of Dianchi in Yunnan Province and Qiongdou in Sichuan Province. After the 3rd century B.C., the ancient Yi extended their activities from the Anning River valley, the Jinsha River, the Dianchi Lake, and the Ailao Mountains to northeastern and southern Yunnan, northwestern Guizhou, and northwestern Guangxi (Zhu et al., 2008 2008 ). Most Yi people are farmers, herders of cattle, sheep and goats, and nomadic hunters. Only about one third of the Yi are literate and most elders are not able to write. The Yi language, which is written in the Yi script, belongs to the Tibeto-Burman language group of the Chinese-Tibetan language family, and includes six dialects. Most Yi are animists, with believes combined with elements of Daoism, shamanism, and fetishism, except for those who have been influenced by Buddhism through the Han culture or converted to Christianism (Guo and Dong, 2000 2000 ). Accounting for 61% of the total Yi population in China, the Yunnan Province ranks the first (Yang et al., 2004 2004 ). The Honghe Hani and Yi Autonomous Prefecture is located in the south of the Yunnan Province of China and borders Vietnam to the south. According to the 2000 national census, Honghe has 973 732 Yi, accounting for 23.57% of the whole population in this area (China, National Bureau of Statistics, 2001 2001 ). Although there are several studies about the polymorphism of HLA genes of the Yi population using low resolution techniques (Chen W.M. et al., 2001 2001 ; Lu et al., 2003a 2003a ; 2003b 2003b ; Wen et al., 2004 2004 ; Xu et al., 2006 2006 ; Chen S. et al., 2007 2007 ), no high-resolution data using HLA class I gene polymorphisms of the Yi people are available now. In the present study we have investigated the distributions of HLA-A and -B alleles and HLA-A-B haplotypes in the Yi ethnic minority in the Honghe Prefecture of the Yunnan Province, southwestern China, using the polymerase chain reaction-sequence-based typing (PCR-SBT) method, and compared these data with ones recorded in other populations in order to enrich the knowledge of HLA polymorphism in Chinese ethnic groups and to better understand the genetic relationships among different populations. This is the first HLA study of the Yi ethnic minority of the Honghe Autonomous Prefecture. Moreover, the distribution of HLA-A-B haplotypes was also investigated for the first time in the Yi in this study. The results from these analyses therefore will provide useful information for blood and bone marrow transplantation, anthropological researches, and forensic science as well as for disease-association studies. Materials and methods 1 Samples and DNA extraction 2 After informed consent, non-coagulated ethylenediaminetetraacetic acid (EDTA) blood samples were obtained from 114 unrelated healthy Yi individuals living in the Honghe Hani and Yi Autonomous Prefecture, Yunnan, China. All participants were randomly chosen among individuals whose ancestors have lived in the region for at least three generations and were self-identified as Yi. All blood samples were stored at −20 °C before DNA extraction. Genomic DNA was extracted using standard techniques (Miller et al., 1988 1988 ). Typing of HLA class I genes 2 All subjects were typed for HLA-A and -B loci. SBT of the HLA-A and -B loci was performed on exons 2 and 3, according to Kurz et al. ( 1999 1999 ) and Pozzi et al. ( 1999 1999 ) with minor modifications. Amplification by polymerase chain reaction (PCR) was accomplished on GeneAmp PCR system 9700 (Applied Biosystems, Foster City, CA, USA). PCR-amplified DNA fragments were purified and sequenced using the ABI PRISM BigDye Terminator Cycle Sequencing Ready Reaction Kits (Applied Biosystems, Foster City, CA, USA) using an ABI 3730XL DNA sequencer (Applied Biosystems, Foster City, CA, USA), according to the manufacturer’s instructions. Sequencing was always performed in forward and reverse directions and processed using Sequencing Analysis, MatchTools and Navigator (MatchTools Allele Identification package, Applied Biosystems, Foster City, CA, USA). This software was used to detect the heterozygous positions within each electropherogram and to assess the typing, based on an alignment of the processed sequence with an updated library of HLA sequences and alleles defined up to October 2007. Ambiguous types were resolved to four digits according to the updated database (IMGT release 2.19.0). Statistical analysis 2 Allele frequencies of HLA-A and -B loci were estimated using SPSS 11.0 software (SPSS Inc., Chicago, Illinois). Haplotype frequencies were calculated from genotype data by expectation maximization (EM) algorithm using Arlequin software package version 3.0 (Laurent Excoffier, CMPG, Zoological Institute, University of Bern, Switzerland) (Excoffier et al., 2005 2005 ). Forensic parameters including observed heterozygosity (Ho), expected heterozygosity (He), power of discrimination (PD), polymorphism information content (PIC), and probability of paternity exclusion (PPE) were computed using the PowerStat version 1.2 spreadsheet (Promega Corporation, USA) as described (Tereba, 1999 1999 ). The exact test of Guo and Thompson ( 1992 1992 ) was used to evaluate deviation from expected Hardy-Weinberg genotypic proportions. Absolute linkage disequilibrium (ALD), maximal linkage disequilibrium (MLD), and relative linkage disequilibrium (RLD) between two alleles at the HLA-A and -B loci were calculated according to Weir ( 1996 1996 ). Analysis of molecular variance (AMOVA) was performed by Arlequin to estimate the population differentiation and to compare HLA allele frequencies between Yi and other populations, and the results are presented as the fixation index. A neighbor-joining (NJ) tree from allele frequencies of HLA-A and -B loci was constructed using the Mega3.1 software package (Center for Evolutionary Functional Genomics, the Biodesign Institute Tempe, AZ, USA). Results 1 Study design 2 Allele frequencies of HLA class I genes were investigated by PCR-SBT in 114 unrelated subjects from the Yi ethnic minority in the Yunnan Province of southwestern China. The Ewens-Watterson homozygosity test showed no evidence for a departure from selective neutrality at HLA-A and -B loci. No significant deviation from the Hardy-Weinberg equilibrium was observed at the HLA-A and HLA-B loci ( P =0.05249 and 0.103721, respectively). Allele distribution 2 In total, we detected 17 HLA-A and 38 HLA-B alleles in the subjects. The allelic distributions are summarized in Table 1 1 . The three most common HLA-A alleles with a frequency greater than 10% were HLA-A*2402 (32.46%), A*1101 (26.32%), and A*0203 (10.09%). The second in rank with frequencies greater than 3% were alleles A*0207, A*0201, and A*3101. At the HLA-B locus, B*4601 had the highest frequency (12.28%), which was followed by B*1525 (10.09%), B*4001 (8.77%), and B*3802 (7.89%). HLA-A AF (%) HLA-B AF (%) HLA-B AF (%) A*0101 0.44 B*0702 1.32 B*3802 7.89 A*0201 6.14 B*0705 0.44 B*3901 2.19 A*0203 10.09 B*1301 3.95 B*4001 8.77 A*0205 0.44 B*1501 7.02 B*4002 1.75 A*0206 1.75 B*1502 7.46 B*4006 0.88 A*0207 7.02 B*1504 0.44 B*4402 0.44 A*0258 0.44 B*1507 1.75 B*4403 1.32 A*0301 0.88 B*1511 0.44 B*4601 12.28 A*1101 26.32 B*1512 2.63 B*4801 1.32 A*1102 0.44 B*1518 0.44 B*5101 3.51 A*2402 32.46 B*1525 10.09 B*5102 0.44 A*2403 2.63 B*1527 0.44 B*5121 0.44 A*2407 2.19 B*1532 0.44 B*5201 4.82 A*2601 2.19 B*2705 0.88 B*5401 0.88 A*2901 0.44 B*2706 0.44 B*5501 0.44 A*3101 3.51 B*3501 3.07 B*5502 5.26 A*3303 2.63 B*3505 1.32 B*5601 2.63 B*3543 0.44 B*5602 0.44 B*3701 0.44 B*5604 0.88 Haplotype frequencies 2 As summarized in Table 2 2 , of the 180 HLA-A-B haplotypes predicted by the EM algorithm, 37 were found at frequencies greater than 0.58%. Two HLA-A-B haplotypes occurred at frequencies greater than 5%, i.e., HLA-A*2402-B*1525 (7.86%) and A*0203-B*3802 (5.64%), followed by A*1101-B*4001 (4.69%), A*0207-B*4601 (4.64%), A*1101-B*5201 (4.37%), and A*2402-B*4601 (4.09%). Table 2 2 also shows the values of ALD, MLD, and RLD, which are commonly used in the evaluation of the linkage disequilibrium (LD) in population genetics. Twenty-seven HLA-A-B haplotypes are common haplotypes with significant linkage disequilibrium (both haplotype frequency and RLD are higher than 1%). HF: haplotype frequency; ALD: absolute linkage disequilibrium; MLD: maximal linkage disequilibrium; RLD: relative linkage disequilibrium HLA-A-B haplotype HF (%) ALD (%) MLD (%) RLD (%) A*2402-B*1525 7.86 4.54 6.77 67.04 A*0203-B*3802 5.64 4.85 7.10 68.30 A*1101-B*4001 4.69 2.38 6.46 36.80 A*0207-B*4601 4.64 3.78 6.16 61.35 A*1101-B*5201 4.37 3.10 3.55 87.31 A*2402-B*4601 4.09 0.05 8.24 0.65 A*1101-B*1502 3.30 1.34 5.49 24.39 A*2402-B*4001 3.06 0.17 5.89 2.89 A*2402-B*1501 2.63 0.32 4.71 6.86 A*1101-B*1301 2.60 1.56 2.91 53.63 A*2402-B*5502 2.41 0.68 3.53 19.32 A*1101-B*5502 2.41 1.03 3.88 26.46 A*2403-B*1512 1.75 1.69 2.56 65.77 A*0201-B*1501 1.71 1.28 5.71 22.38 A*0201-B*4601 1.60 0.84 5.39 15.63 A*1101-B*5101 1.58 0.66 2.59 25.36 A*0201-B*1502 1.36 0.90 5.68 15.91 A*2402-B*1301 1.35 0.05 2.65 1.890 A*2402-B*4801 1.32 0.88 0.88 100.00 A*1101-B*0702 1.32 0.97 0.97 100.00 A*3303-B*4403 1.32 1.28 1.28 100.00 A*1101-B*4002 1.32 0.85 1.29 66.07 A*2402-B*1507 1.32 0.74 1.18 62.75 A*2402-B*3501 1.30 0.29 2.06 13.90 A*0203-B*3901 1.27 1.04 1.97 52.95 A*0203-B*1525 1.25 0.23 9.07 2.58 A*0203-B*4001 1.03 0.14 7.89 1.83 A*0207-B*5604 0.88 0.82 0.82 100.00 A*2601-B*2705 0.88 0.86 0.86 100.00 A*0206-B*4006 0.88 0.86 0.86 100.00 A*0206-B*1512 0.88 0.83 1.71 48.65 A*2601-B*5601 0.88 0.82 2.14 38.38 A*2407-B*3501 0.88 0.81 2.13 38.10 A*3101-B*5601 0.88 0.78 2.54 30.91 A*3101-B*1501 0.88 0.63 3.26 19.34 A*2402-B*5401 0.72 0.43 0.59 73.11 A*0207-B*3802 0.62 0.07 6.46 1.09 Forensic parameters 2 The values of common forensic parameters of HLA-A and -B loci in the Yi population are listed in Table 3 3 . The Ho, He, PD, PIC, and PPE were 0.7997, 0.7456, 0.9289, 0.7762, 0.5023 for HLA-A and 0.9382, 0.8596, 0.9846, 0.9349, 0.7140 for HLA-B, respectively. Ho: observed heterozygosity; He: expected heterozygosity; PD: power of discrimination; PIC: polymorphism information content; PPE: probability of paternity exclusion Locus Ho He PD PIC PPE HLA-A 0.7997 0.7456 0.9289 0.7762 0.5023 HLA-B 0.9382 0.8596 0.9846 0.9349 0.7140 Phylogenetic tree analysis 2 As shown in Fig. 1 1 , an NJ phylogenetic tree was constructed using the allele frequencies of HLA-A and -B loci of 28 representative populations all over the world. Two main clusters were obtained: African and Asian/European. Located in the southeastern Asian cluster, the Yi population was first grouped with Tujia ethnic group and followed by Jinuo, Wa, Shui, Miao and Bouyei ethnic minorities living in China, as well as Vietnamese and Javanese groups. Subsequently, they were joined by the southern Chinese groups, including the Taiwanese, Hong Kong Chinese, Singapore Chinese, and southern Canton Chinese. Compared to other ethnic minorities like Jinuo, Wa, Shui, Miao, and Bouyei, the genetic structure of the Yi is the closest to the northeastern Asian populations. Discussion 1 The HLA is the most polymorphic genetic system of human genome, consisting of a number of closely linked genes (Marsh et al., 2002 2002 ). The distributions of HLA alleles and haplotypes have been widely used in the investigation of the origin and migration of human populations, transplantation matching, forensic sciences, and disease-association studies (Li et al., 2009 2009 ; Wang et al., 2009 2009 ). The technology of HLA typing has significantly evolved since the PCR method was introduced by Mullis and Faloona ( 1987 1987 ). Nowadays, there are several kinds of PCR-based HLA typing being used: DNA amplification with sequence specific primers (SSP) (Zetterquist and Olerup, 1992 1992 ), sequence-specific oligonucleotide probes (SSOP) (Petersdorf et al., 1991 1991 ), single-stranded conformation polymorphism (SSCP) (Lo et al., 1992 1992 ), sequence-based typing (SBT) (Santamaria et al., 1993 1993 ), and DNA chip technology (Jiang et al., 2006 2006 ). These HLA typing techniques have much higher accuracy as well as reliability than the classical serologic typing method, and also facilitate the standardization of HLA typing processes. Among all the PCR-based HLA typing, SBT is considered the gold standard (Danzer et al., 2007 2007 ) and the most comprehensive method for characterizing MHC gene polymorphisms (Takeshima et al., 2009 2009 ). Moreover, new HLA alleles can be easily identified by SBT and confirmed after cloning (Scheltinga et al., 1997 1997 ). We found that there were significant differences between the studied Yi population living in Yunnan and the one living in Sichuan at HLA-A ( P =0.037) (Lu et al., 2003a 2003a ) and HLA-B ( P < 0.001) (Xu et al., 2006 2006 ). Studies have shown that allelic distributions in the HLA-A and -B loci differ from the members of the Chinese Yi ethnic group living in different geographic areas. In Asians, HLA-A*1101, A*2402, and A*3303 were found at significantly higher frequencies than those observed in other populations (Cao et al., 2001 2001 ). As the predominant HLA-A allele in the Yi population, A*2402 (32.46%) is also the most common one of other populations such as the Tibetan (Chen et al., 2006 2006 ), Mongolian (Hong et al., 2007 2007 ), Japanese (Saito et al., 2000 2000 ), and Korean groups (Lee et al., 2005 2005 ) in northern Asia and interestingly, also in north native Americans (Cao et al., 2001 2001 ). A*1101, the second most common HLA-A allele in the Yi (26.32%), has the highest prevalence in Southwest Asia (Chen et al., 2007 2007 ; Ogata et al., 2007 2007 ; Hoa et al., 2008 2008 ; Shi L. et al., 2008 2008 ) and in many Chinese population groups (20.23%–28.70%) (Middleton et al., 2004 2004 ; Yang et al., 2006 2006 ; Trachtenberg et al., 2007 2007 ; Wen et al., 2008 2008 ). A*1101 and A*2402 together accounting for almost 60% of all HLA-A alleles were observed in the Yi population, whereas A*3303 had a relatively low frequency (2.63%) than that in other Asian populations such as the Korean (16.49%) (Lee et al., 2005 2005 ), Wa ethnic minority living in China (16.10%) (Shi L. et al., 2008 2008 ), Javanese (15.61%) (Yuliwulandari et al., 2009 2009 ), Vietnamese (11.50%) (Hoa et al., 2008 2008 ), and other Chinese groups (6.23%–11.70%) (Middleton et al., 2004 2004 ; Yang et al., 2006 2006 ; Trachtenberg et al., 2007 2007 ; Wen et al., 2008 2008 ). These observations suggest that the Yi people have some characteristics of northeastern Asian groups in terms of HLA-A allelic distribution. In the Yi people, B*4601 had the highest prevalence (12.28%) at the HLA-B locus. In a disease association study between the HLA allele distribution and the epidemic of severe acute respiratory syndrome (SARS), Lin et al. ( 2003 2003 ) showed that HLA-B*4601 was a possible susceptibility HLA allele for the infection of SARS. In previous studies, HLA-B*4601 was also reported to occur at high frequencies in Southeast Asian populations such as the Vietnamese population (11.50%) (Hoa et al., 2008 2008 ) and in southern Chinese groups such as in the Taiwanese (Wen et al., 2008 2008 ), Hong Kong Chinese (Middleton et al., 2004 2004 ), Singapore Chinese (Middleton et al., 2004 2004 ), and southern Canton Chinese groups (Trachtenberg et al., 2007 2007 ) (11.90%–16.30%), where the highest occurrence of SARS cases of the world was found. By contrast, B*4601 is rarely seen in European populations (0–0.9%) (Cao et al., 2001 2001 ; Spínola et al., 2005a 2005a ; 2005b 2005b ; Dunne et al., 2008 2008 ; Mack et al., 2009 2009 ), and never seen in African populations (Cao et al., 2001 2001 ; 2004 2004 ) or in native Americans (Cao et al., 2001 2001 ), where very few cases of SARS infection were reported. HLA-B*1502, a common HLA-B allele in the Yi population (7.46%) and in Southeast Asia, was reported to have a strong association with cutaneous adverse drug reactions, including carbamazepine (CBZ)-induced Stevens-Johnson syndrome (SJS) and toxic epidermal necrolysis (TEN) in Han Chinese and many other Asian populations (Chung et al., 2004 2004 ; Hung et al., 2006 2006 ; Lonjou et al., 2006 2006 ; Man et al., 2007 2007 ). So far, there have been no epidemiologic studies of SARS, CBZ-SJS, or TEN conducted in the Yi population. As one of the most important characteristics of the HLA system, the LD between the HLA-A and -B loci in the Yi population was measured through calculating ALD, MLD, and RLD. Six HLA-A-B haplotypes, including A*2402-B*4801, A*1101-B*0702, A*3303-B*4403, A*0207-B*5604, A*2601-B*2705, and A*0206-B*4006, had the most significant LD ( RLD =1), which indicates that there is very little chance of the recombination between the A and B alleles for these haplotypes in the Yi population. The haplotypes in this study were estimated using EM algorithm, but the haplotypes presented were not based on family data; therefore, we need to acknowledge the potential errors inherent to haplotype estimation methods (Single et al., 2002 2002 ). In the NJ-constructed phylogenic tree, the Yi population has the closest genetic distance to the Tujia ethnic group that was sampled in the Hubei Province of China. The Tujia ethnic group may trace back their history to the ancient Ba people who occupied the area around modern-day Chongqing in southwestern China some 2500 years ago ( http://en.wikipedia.org/wiki/Tujia http://en.wikipedia.org/wiki/Tujia ). The close genetic relationship between Yi and Tujia is also supported by the origin of their languages, which have grammatical and phonological similarities ( http://www.brassett.org.uk/tujia/elng/elintro.html http://www.brassett.org.uk/tujia/elng/elintro.html ) belonging to the Tibeto-Burman language family (often considered a subgroup of the Sino-Tibetan language family) (Shi M. et al., 2008 2008 ), and are spoken in various central, eastern, southern, and southeastern Asian countries. In a previous study of genetic structures in Chinese populations, it was shown that although the Tujia people belong to southern populations of China, they also contain certain lineages that are seen in northern populations. It was also shown that the Yi population was the first to be grouped with the Tujia group, followed by the Bouyei and Wa populations (Du et al., 1997 1997 ), which is in accordance with the observation in this study. Some scholars consider that the Yi descended from the ancient Qiang people of today’s western China, who are also said to be the ancestors of the Tibetan, Naxi, and Qiang peoples. They migrated from southeastern Tibet through Sichuan into Yunnan, where their largest populations can be found today. This may explain that the northeastern Asian groups are closer in the phylogenetic tree to the Yi population (Fig. 1 1 ) than other southeastern Asian ethnic groups located in adjacent regions to the Yi, such as Miao, Bouyei, Shui, Jinuo, and Wa, which have different origins. In a previous study, it was concluded that the Yi originated from the north of the Yangtze River where a significant fraction of them still live, but they at present belong to the southern group (Du et al., 1997 1997 ). As a highly polymorphic genetic marker system, HLA has been applied to forensic science for a long time (Mayr, 1988 1988 ; Lee et al., 1994 1994 ) and is considered a useful tool for personal identification and paternity testing (Allen et al., 1993 1993 ; Li et al., 2004 2004 ). The polymorphisms of genetic markers can be evaluated by some forensic parameters such as Ho, He, PD, PIC, and PPE. Using these parameters, gene heterozygosity can objectively reflect the genetic variation level in a population. Polymorphism information content is often used to measure the indicative strength of genetic markers for linkage studies. A locus is considered highly polymorphic when its PIC is higher than 0.5. Also, PD and PPE are indicators for discrimination capability of a genetic marker and the PD and PPE values of a marker with high polymorphism are normally higher than 0.8 and 0.5, respectively. In our study, the PIC values of HLA-A and -B loci in the Yi population were both higher than 0.5 (0.7762 and 0.9349, respectively) and the PD and PPE at both HLA-A and -B loci were higher than the cutoff values mentioned above. Therefore, the HLA-A and -B polymorphism data obtained from the Yi people were valuable for forensic science research. In conclusion, the HLA polymorphism data presented in this study indicate that the Yi population in Honghe, Yunnan, China, basically belongs to groups of southeastern Asian origin, but at the same time they share some characteristics with northeastern Asian group in terms of the distribution of allele frequencies. This study adds to our understanding of HLA polymorphisms in the ethnic groups in Asia. In addition, these results may provide useful information for blood and bone marrow transplantation, forensic science, and disease-association studies. However, it will be necessary to perform further HLA analysis on class II and other class I loci in order to describe the complete profiles of the Yi population. References M Allen , T Saldeen , U Pettersson , U Gyllensten . Genetic typing of HLA class II genes in Swedish populations: application to forensic analysis . Journal of Forensic Science , 1993 . 38 ( 3 ): 554 - 570 . . Genetic typing of HLA class II genes in Swedish populations: application to forensic analysis K Cao , J Hollenbach , X Shi , W Shi , M Chopek , MA Fernández-Viña . Analysis of the frequencies of HLA-A, B, and C alleles and haplotypes in the five major ethnic groups of the United States reveals high levels of diversity in these loci and contrasting distribution patterns in these populations . Human Immunology , 2001 . 62 ( 9 ): 1009 - 1030 . 11543903 DOI: 10.1016/S0198-8859(01)00298-1 http://doi.org/10.1016/S0198-8859(01)00298-1 . Analysis of the frequencies of HLA-A, B, and C alleles and haplotypes in the five major ethnic groups of the United States reveals high levels of diversity in these loci and contrasting distribution patterns in these populations K Cao , AM Moormann , KE Lyke , C Masaberg , OP Sumba , OK Doumbo , D Koech , A Lancaster , M Nelson , D Meyer , 等 . Differentiation between African populations is evidenced by the diversity of alleles and haplotypes of HLA class I loci . Tissue Antigens , 2004 . 63 ( 4 ): 293 - 325 . 15009803 DOI: 10.1111/j.0001-2815.2004.00192.x http://doi.org/10.1111/j.0001-2815.2004.00192.x . Differentiation between African populations is evidenced by the diversity of alleles and haplotypes of HLA class I loci S Chen , W Hong , H Shao , Y Fu , X Liu , D Chen , A Xu . Allelic distribution of HLA class I genes in the Tibetan ethnic population of China . International Journal of Immunogenetics , 2006 . 33 ( 6 ): 439 - 445 . 17117954 DOI: 10.1111/j.1744-313X.2006.00637.x http://doi.org/10.1111/j.1744-313X.2006.00637.x . Allelic distribution of HLA class I genes in the Tibetan ethnic population of China S Chen , X Ren , Y Liu , Q Hu , W Hong , A Xu . Human leukocyte antigen class I polymorphism in Miao, Bouyei, and Shui ethnic minorities of Guizhou, China . Human Immunology , 2007 . 68 ( 11 ): 928 - 933 . 18082574 DOI: 10.1016/j.humimm.2007.09.006 http://doi.org/10.1016/j.humimm.2007.09.006 . Human leukocyte antigen class I polymorphism in Miao, Bouyei, and Shui ethnic minorities of Guizhou, China WM Chen , HL Huang , ZH Liu , H Tao , DJ Pan , JH Lin , AL Xu . Analysis of HLA-DQB1 polymorphism by PCR-SSO in Yichu of Yunnan Province . Yi Chuan Xue Bao , 2001 . 28 ( 2 ): 107 - 114 . (in Chinese) 11233253 . Analysis of HLA-DQB1 polymorphism by PCR-SSO in Yichu of Yunnan Province China, National Bureau of Statistics Chinese Statistical Yearbook 2000 , 2001 . : Beijing, China China Statistics Press , 415 - 416 . (in Chinese) . Chinese Statistical Yearbook 2000 WH Chung , SI Hung , HS Hong , MS Hsih , LC Yang , HC Ho , JY Wu , YT Chen . Medical genetics: a marker for Stevens-Johnson syndrome . Nature , 2004 . 428 ( 6982 ): 486 15057820 DOI: 10.1038/428486a http://doi.org/10.1038/428486a . Medical genetics: a marker for Stevens-Johnson syndrome M Danzer , H Polin , J Pröll , K Hofer , I Fae , GF Fischer , C Gabriel . High-throughput sequence-based typing strategy for HLA-DRB1 based on real-time polymerase chain reaction . Human Immunology , 2007 . 68 ( 11 ): 915 - 917 . 18082572 DOI: 10.1016/j.humimm.2007.10.005 http://doi.org/10.1016/j.humimm.2007.10.005 . High-throughput sequence-based typing strategy for HLA-DRB1 based on real-time polymerase chain reaction R Du , C Xiao , LL Cavalli-sforza . Genetic distances between Chinese populations calculated on gene frequencies of 38 loci . Science in China Series C, Life Sciences , 1997 . 40 ( 6 ): 613 - 621 . (in Chinese) DOI: 10.1007/BF02882691 http://doi.org/10.1007/BF02882691 . Genetic distances between Chinese populations calculated on gene frequencies of 38 loci C Dunne , J Crowley , R Hagan , G Rooney , E Lawlor . HLA-A, B, Cw, DRB1, DQB1 and DPB1 alleles and haplotypes in the genetically homogenous Irish population . International Journal of Immunogenetics , 2008 . 35 ( 4-5 ): 295 - 302 . 18976432 DOI: 10.1111/j.1744-313X.2008.00779.x http://doi.org/10.1111/j.1744-313X.2008.00779.x . HLA-A, B, Cw, DRB1, DQB1 and DPB1 alleles and haplotypes in the genetically homogenous Irish population HA Erlich , G Opelz , J Hansen . HLA DNA typing and transplantation . Immunity , 2001 . 14 ( 4 ): 347 - 356 . 11336680 DOI: 10.1016/S1074-7613(01)00115-7 http://doi.org/10.1016/S1074-7613(01)00115-7 . HLA DNA typing and transplantation L Excoffier , G Laval , S Schneider . Arlequin (version 3.0): an integrated software package for population genetics data analysis . Evolutionary Bioinformatics Online , 2005 . 1 47 - 50 . 19325852 . Arlequin (version 3.0): an integrated software package for population genetics data analysis DL Guo , JZ Dong . Summarization of Chinese Nationalities (Zhonghua Mingzu Zhishi Tonglan) , 2000 . : Kunming, China Kunming Education Publishing House , 86 - 94 . (in Chinese) . Summarization of Chinese Nationalities (Zhonghua Mingzu Zhishi Tonglan) SW Guo , EA Thompson . Performing the exact test of Hardy-Weinberg proportion for multiple alleles . Biometrics , 1992 . 48 ( 2 ): 361 - 372 . 1637966 DOI: 10.2307/2532296 http://doi.org/10.2307/2532296 . Performing the exact test of Hardy-Weinberg proportion for multiple alleles BK Hoa , NT Hang , K Kashiwase , J Ohashi , LT Lien , T Horie , J Shojima , M Hijikata , S Sakurada , M Satake , 等 . HLA-A, -B, -C, -DRB1 and -DQB1 alleles and haplotypes in the Kinh population in Vietnam . Tissue Antigens , 2008 . 71 ( 2 ): 127 - 134 . 18086267 . HLA-A, -B, -C, -DRB1 and -DQB1 alleles and haplotypes in the Kinh population in Vietnam W Hong , S Chen , H Shao , Y Fu , Z Hu , A Xu . HLA class I polymorphism in Mongolian and Hui ethnic groups from Northern China . Human Immunology , 2007 . 68 ( 5 ): 439 - 448 . 17462512 DOI: 10.1016/j.humimm.2007.01.020 http://doi.org/10.1016/j.humimm.2007.01.020 . HLA class I polymorphism in Mongolian and Hui ethnic groups from Northern China SP Hu , JA Luan , B Li , JX Chen , KL Cai , LQ Huang , XY Xu . Genetic link between Chaoshan and other Chinese Han populations: evidence from HLA-A and HLA-B allele frequency distribution . American Journal of Physical Anthropology , 2007 . 132 ( 1 ): 140 - 150 . 16883565 DOI: 10.1002/ajpa.20460 http://doi.org/10.1002/ajpa.20460 . Genetic link between Chaoshan and other Chinese Han populations: evidence from HLA-A and HLA-B allele frequency distribution SI Hung , WH Chung , SH Jee , WC Chen , YT Chang , WR Lee , SL Hu , MT Wu , GS Chen , TW Wong , 等 . Genetic susceptibility to carbamazepine-induced cutaneous adverse drug reactions . Pharmaco-genetics and Genomics , 2006 . 16 ( 4 ): 297 - 306 . DOI: 10.1097/01.fpc.0000199500.46842.4a http://doi.org/10.1097/01.fpc.0000199500.46842.4a . Genetic susceptibility to carbamazepine-induced cutaneous adverse drug reactions B Jiang , Y Li , H Wu , XM He , CT Li , L Li , R Tang , Y Xie , YM Mao . Application of HLA-DRB1 genotyping by oligonucleotide micro-array technology in forensic medicine . Forensic Science International , 2006 . 162 ( 1-3 ): 66 - 73 . 16884876 DOI: 10.1016/j.forsciint.2006.06.041 http://doi.org/10.1016/j.forsciint.2006.06.041 . Application of HLA-DRB1 genotyping by oligonucleotide micro-array technology in forensic medicine B Kurz , I Steiert , G Heuchert , CA Müller . New high resolution typing strategy for HLA-A locus alleles based on dye terminator sequencing of haplotypic group-specific PCR-amplicons of exon 2 and exon 3 . Tissue Antigens , 1999 . 53 ( 1 ): 81 - 96 . 10082434 DOI: 10.1034/j.1399-0039.1999.530109.x http://doi.org/10.1034/j.1399-0039.1999.530109.x . New high resolution typing strategy for HLA-A locus alleles based on dye terminator sequencing of haplotypic group-specific PCR-amplicons of exon 2 and exon 3 HC Lee , C Ladd , MT Bourke , E Pagliaro , F Tirnady . DNA typing in forensic science. I. Theory and background . The American Journal of Forensic Medicine and Pathology , 1994 . 15 ( 4 ): 269 - 282 . 7879769 DOI: 10.1097/00000433-199412000-00001 http://doi.org/10.1097/00000433-199412000-00001 . DNA typing in forensic science. I. Theory and background KW Lee , DH Oh , C Lee , SY Yang . Allelic and haplotypic diversity of HLA-A, -B, -C, -DRB1, and -DQB1 genes in the Korean population . Tissue Antigens , 2005 . 65 ( 5 ): 437 - 447 . 15853898 DOI: 10.1111/j.1399-0039.2005.00386.x http://doi.org/10.1111/j.1399-0039.2005.00386.x . Allelic and haplotypic diversity of HLA-A, -B, -C, -DRB1, and -DQB1 genes in the Korean population L Li , CT Li , Y Liu . Research on the application feasibility of HLA-DRB1 genotyping for forensic identification by oligonucleotide chip . Fa Yi Xue Za Zhi , 2004 . 20 ( 2 ): 81 - 84 . 81 (in Chinese) 15311520 . Research on the application feasibility of HLA-DRB1 genotyping for forensic identification by oligonucleotide chip Y Li , H Yan , WJ Xue , PX Tian , XM Ding , XM Pan , XS Feng , XH Tian , HL Xiang , J Hou . Allograft rejection-related gene expression in the endothelial cells of renal transplantation recipients after cytomegalovirus infection . Journal of Zhejiang University-SCIENCE B , 2009 . 10 ( 11 ): 820 - 828 . 19882756 DOI: 10.1631/jzus.B0920115 http://doi.org/10.1631/jzus.B0920115 . Allograft rejection-related gene expression in the endothelial cells of renal transplantation recipients after cytomegalovirus infection M Lin , HK Tseng , JA Trejaut , HL Lee , JH Loo , CC Chu , PJ Chen , YW Su , KH Lim , ZU Tsai , 等 . Association of HLA class I with severe acute respiratory syndrome coronavirus infection . BMC Medical Genetics , 2003 . 4 ( 1 ): 9 12969506 DOI: 10.1186/1471-2350-4-9 http://doi.org/10.1186/1471-2350-4-9 . Association of HLA class I with severe acute respiratory syndrome coronavirus infection YM Lo , P Patel , WZ Mehal , KA Fleming , JI Bell , JS Wainscoat . Analysis of complex genetic systems by ARMS-SSCP: application to HLA genotyping . Nucleic Acids Research , 1992 . 20 ( 5 ): 1005 - 1009 . 1549460 DOI: 10.1093/nar/20.5.1005 http://doi.org/10.1093/nar/20.5.1005 . Analysis of complex genetic systems by ARMS-SSCP: application to HLA genotyping C Lonjou , L Thomas , N Borot , N Ledger , C de Toma , H LeLouet , E Graf , M Schumacher , A Hovnanian , M Mockenhaupt , 等 . A marker for Stevens-Johnson syndrome …: ethnicity matters . The Pharmacogenomics Journal , 2006 . 6 ( 4 ): 265 - 268 . 16415921 DOI: 10.1038/sj.tpj.6500356 http://doi.org/10.1038/sj.tpj.6500356 . A marker for Stevens-Johnson syndrome …: ethnicity matters XZ Lu , KX Hong , GM Qin , CX Li , JH Zhu , YM Shao . DNA typing of HLA-A locus in Chinese Yi ethnic group by PCR-SSP . Immunological Journal , 2003 . 19 ( 6 ): 2 (in Chinese) . DNA typing of HLA-A locus in Chinese Yi ethnic group by PCR-SSP XZ Lu , KX Hong , GM Qin , JP Chen , YH Ruan , CH Li , JH Zhu , YM Shao . Genotyping of HLA-Cw locus in Chinese Yi ethnic group by PCR-SSP . Zhonghua Shi Yan He Lin Chuang Bing Du Xue Za Zhi , 2003 . 17 ( 1 ): 62 - 65 . (in Chinese) 12870022 . Genotyping of HLA-Cw locus in Chinese Yi ethnic group by PCR-SSP SJ Mack , B Tu , A Lazaro , R Yang , AK Lancaster , K Cao , J Ng , CK Hurley . HLA-A, -B, -C, and -DRB1 allele and haplotype frequencies distinguish eastern European Americans from the general European American population . Tissue Antigens , 2009 . 73 ( 1 ): 17 - 32 . 19000140 DOI: 10.1111/j.1399-0039.2008.01151.x http://doi.org/10.1111/j.1399-0039.2008.01151.x . HLA-A, -B, -C, and -DRB1 allele and haplotype frequencies distinguish eastern European Americans from the general European American population CB Man , P Kwan , L Baum , E Yu , KM Lau , AS Cheng , MH Ng . Association between HLA-B*1502 allele and antiepileptic drug-induced cutaneous reactions in Han Chinese . Epilepsia , 2007 . 48 ( 5 ): 1015 - 1018 . 17509004 DOI: 10.1111/j.1528-1167.2007.01022.x http://doi.org/10.1111/j.1528-1167.2007.01022.x . Association between HLA-B*1502 allele and antiepileptic drug-induced cutaneous reactions in Han Chinese SG Marsh , ED Albert , WF Bodmer , RE Bontrop , B Dupont , HA Erlich , DE Geraghty , JA Hansen , B Mach , WR Mayr , 等 . Nomenclature for factors of the HLA system, 2002 . European Journal of Immunogenetics , 2002 . 29 ( 6 ): 463 - 515 . 12437610 DOI: 10.1046/j.1365-2370.2002.00359.x http://doi.org/10.1046/j.1365-2370.2002.00359.x . Nomenclature for factors of the HLA system, 2002 WR Mayr . The detection of HLA antigens in bloodstains . Zeitschrift für Rechtsmedizin Journal of Legal Medicine , 1988 . 101 ( 4 ): 209 - 217 . . The detection of HLA antigens in bloodstains D Middleton , BR Hawkins , F Williams , A Meenagh , J Moscoso , J Zamora , A Arnaiz-Villena . HLA class I allele distribution of a Hong Kong Chinese population based on high-resolution PCR-SSOP typing . Tissue Antigens , 2004 . 63 ( 6 ): 555 - 561 . 15140031 DOI: 10.1111/j.0001-2815.2004.00234.x http://doi.org/10.1111/j.0001-2815.2004.00234.x . HLA class I allele distribution of a Hong Kong Chinese population based on high-resolution PCR-SSOP typing SA Miller , DD Dykes , HF Polesky . A simple salting out procedure for extracting DNA from human nucleated cells . Nucleic Acids Research , 1988 . 16 ( 3 ): 1215 3344216 DOI: 10.1093/nar/16.3.1215 http://doi.org/10.1093/nar/16.3.1215 . A simple salting out procedure for extracting DNA from human nucleated cells KB Mullis , FA Faloona . Specific synthesis of DNA in vitro via a polymerase-catalyzed chain reaction . Methods in Enzymology , 1987 . 155 335 - 350 . 3431465 DOI: 10.1016/0076-6879(87)55023-6 http://doi.org/10.1016/0076-6879(87)55023-6 . Specific synthesis of DNA in vitro via a polymerase-catalyzed chain reaction S Ogata , L Shi , M Matsushita , L Yu , XQ Huang , L Shi , H Sun , J Ohashi , M Muramatsu , K Tokunaga , 等 . Polymorphisms of human leucocyte antigen genes in Maonan people in China . Tissue Antigens , 2007 . 69 ( 2 ): 154 - 160 . 17257318 DOI: 10.1111/j.1399-0039.2006.00698.x http://doi.org/10.1111/j.1399-0039.2006.00698.x . Polymorphisms of human leucocyte antigen genes in Maonan people in China EW Petersdorf , AG Smith , AM Haase , PJ Martin , JA Hansen . Polymorphism of HLA-DRw52-associated DRB1 genes as defined by sequence-specific oligonucleotide probe hybridization and sequencing . Tissue Antigens , 1991 . 38 ( 4 ): 169 - 177 . 1801307 DOI: 10.1111/j.1399-0039.1991.tb01891.x http://doi.org/10.1111/j.1399-0039.1991.tb01891.x . Polymorphism of HLA-DRw52-associated DRB1 genes as defined by sequence-specific oligonucleotide probe hybridization and sequencing S Pozzi , A Longo , GB Ferrara . HLA-B locus sequence-based typing . Tissue Antigens , 1999 . 53 ( 3 ): 275 - 281 . 10203021 DOI: 10.1034/j.1399-0039.1999.530308.x http://doi.org/10.1034/j.1399-0039.1999.530308.x . HLA-B locus sequence-based typing S Saito , S Ota , E Yamada , H Inoko , M Ota . Allele frequencies and haplotypic associations defined by allelic DNA typing at HLA class I and class II loci in the Japanese population . Tissue Antigens , 2000 . 56 ( 6 ): 522 - 529 . 11169242 DOI: 10.1034/j.1399-0039.2000.560606.x http://doi.org/10.1034/j.1399-0039.2000.560606.x . Allele frequencies and haplotypic associations defined by allelic DNA typing at HLA class I and class II loci in the Japanese population P Santamaria , AL Lindstrom , MT Boyce-Jacino , SH Myster , JJ Barbosa , AJ Faras , SS Rich . HLA class I sequence-based typing . Human Immunology , 1993 . 37 ( 1 ): 39 - 50 . 8376187 DOI: 10.1016/0198-8859(93)90141-M http://doi.org/10.1016/0198-8859(93)90141-M . HLA class I sequence-based typing SA Scheltinga , LA Johnston-Dow , CB White , AW van der Zwan , JE Bakema , EH Rozemuller , JG van den Tweel , MN Kronick , MG Tilanus . A generic sequencing based typing approach for the identification of HLA-A diversity . Human Immunology , 1997 . 57 ( 2 ): 120 - 128 . 9438203 DOI: 10.1016/S0198-8859(97)00204-8 http://doi.org/10.1016/S0198-8859(97)00204-8 . A generic sequencing based typing approach for the identification of HLA-A diversity CM Shen , BF Zhu , ML Liu , SB Li . Genetic polymorphisms at HLA-A, -B, and -DRB1 loci in Han population of Xi’an city in China . Croatian Medical Journal , 2008 . 49 ( 4 ): 476 - 482 . 18716994 DOI: 10.3325/cmj.2008.4.476 http://doi.org/10.3325/cmj.2008.4.476 . Genetic polymorphisms at HLA-A, -B, and -DRB1 loci in Han population of Xi’an city in China L Shi , S Ogata , JK Yu , J Ohashi , L Yu , L Shi , H Sun , K Lin , XQ Huang , M Matsushita , 等 . Distribution of HLA alleles and haplotypes in Jinuo and Wa populations in Southwest China . Human Immunology , 2008 . 69 ( 1 ): 58 - 65 . 18295677 DOI: 10.1016/j.humimm.2007.11.007 http://doi.org/10.1016/j.humimm.2007.11.007 . Distribution of HLA alleles and haplotypes in Jinuo and Wa populations in Southwest China M Shi , R Bai , L Wan , X Yu , L Chang . Population genetics for Y-chromosomal STRs haplotypes of Chinese Tujia ethnic group . Forensic Science International Genetics , 2008 . 2 ( 4 ): e65 - e68 . 19083831 DOI: 10.1016/j.fsigen.2007.12.010 http://doi.org/10.1016/j.fsigen.2007.12.010 . Population genetics for Y-chromosomal STRs haplotypes of Chinese Tujia ethnic group RM Single , D Meyer , JA Hollenbach , MP Nelson , JA Noble , HA Erlich , G Thomson . Haplotype frequency estimation in patient populations: the effect of departures from Hardy-Weinberg proportions and collapsing over a locus in the HLA region . Genetic Epidemiology , 2002 . 22 ( 2 ): 186 - 195 . 11788963 DOI: 10.1002/gepi.0163 http://doi.org/10.1002/gepi.0163 . Haplotype frequency estimation in patient populations: the effect of departures from Hardy-Weinberg proportions and collapsing over a locus in the HLA region H Spínola , A Brehm , B Bettencourt , D Middleton , J Bruges-Armas . HLA class I and II polymorphisms in Azores show different settlements in Oriental and Central islands . Tissue Antigens , 2005 . 66 ( 3 ): 217 - 230 . 16101833 DOI: 10.1111/j.1399-0039.2005.00471.x http://doi.org/10.1111/j.1399-0039.2005.00471.x . HLA class I and II polymorphisms in Azores show different settlements in Oriental and Central islands H Spínola , D Middleton , A Brehm . HLA genes in Portugal inferred from sequence-based typing: in the crossroad between Europe and Africa . Tissue Antigens , 2005 . 66 ( 1 ): 26 - 36 . 15982254 DOI: 10.1111/j.1399-0039.2005.00430.x http://doi.org/10.1111/j.1399-0039.2005.00430.x . HLA genes in Portugal inferred from sequence-based typing: in the crossroad between Europe and Africa SN Takeshima , Y Matsumoto , Y Aida . Short communication: establishment of a new polymerase chain reaction-sequence-based typing method for genotyping cattle major histocompatibility complex class II DRB3 . Journal of Dairy Science , 2009 . 92 ( 6 ): 2965 - 2970 . 19448029 DOI: 10.3168/jds.2008-1999 http://doi.org/10.3168/jds.2008-1999 . Short communication: establishment of a new polymerase chain reaction-sequence-based typing method for genotyping cattle major histocompatibility complex class II DRB3 A Tereba . Tools for analysis of population statistics . Profiles DNA , 1999 . 2 3 . Tools for analysis of population statistics E Trachtenberg , M Vinson , E Hayes , YM Hsu , K Houtchens , H Erlich , W Klitz , Y Hsia , J Hollenbach . HLA class I (A, B, C) and class II (DRB1, DQA1, DQB1, DPB1) alleles and haplotypes in the Han from southern China . Tissue Antigens , 2007 . 70 ( 6 ): 455 - 463 . 17900288 DOI: 10.1111/j.1399-0039.2007.00932.x http://doi.org/10.1111/j.1399-0039.2007.00932.x . HLA class I (A, B, C) and class II (DRB1, DQA1, DQB1, DPB1) alleles and haplotypes in the Han from southern China HF Wang , YZ Cheng , HP Wang , ZM Chen , JY Lou , J Jin . CD19-positive acute myeloblastic leukemia with trisomy 21 as a sole acquired karyotypic abnormality . Journal of Zhejiang University-SCIENCE B , 2009 . 10 ( 11 ): 833 - 838 . 19882758 DOI: 10.1631/jzus.B0820362 http://doi.org/10.1631/jzus.B0820362 . CD19-positive acute myeloblastic leukemia with trisomy 21 as a sole acquired karyotypic abnormality BS Weir . Genetic Data Analysis II: Methods for Discrete Population Genetic Data , 1996 . : Sunderland, Massachusetts, United States Sinauer Associates Inc , 204 - 206 . . Genetic Data Analysis II: Methods for Discrete Population Genetic Data G Wen , Y Huang , P Hao , Q Qi , H Li , L Zhou , L Zhou , L Yu . Analysis of HLA-DRB1, DQB1 allele polymorphism in the Kunming Yi nationality population . Zhonghua Yi Xue Yi Chuan Xue Za Zhi , 2004 . 21 ( 5 ): 522 - 523 . (in Chinese) 15476187 . Analysis of HLA-DRB1, DQB1 allele polymorphism in the Kunming Yi nationality population SH Wen , MJ Lai , KL Yang . Human leukocyte antigen-A, -B, and -DRB1 haplotypes of cord blood units in the Tzu Chi Taiwan Cord Blood Bank . Human Immunology , 2008 . 69 ( 7 ): 430 - 436 . 18582515 DOI: 10.1016/j.humimm.2008.05.012 http://doi.org/10.1016/j.humimm.2008.05.012 . Human leukocyte antigen-A, -B, and -DRB1 haplotypes of cord blood units in the Tzu Chi Taiwan Cord Blood Bank MY Xu , KX Hong , J Ma , XL Deng , J Li , H Peng , YH Ruan , GM Qin , YZ Zhang , H Xing , 等 . Analysis of HLA-B locus gene polymorphism in Sichuan Yi ethnic group and Xinjiang Uygur ethnic group . Yi Chuan , 2006 . 28 ( 8 ): 913 - 917 . (in Chinese) 16870575 . Analysis of HLA-B locus gene polymorphism in Sichuan Yi ethnic group and Xinjiang Uygur ethnic group G Yang , YJ Deng , SN Hu , DY Wu , SB Li , J Zhu , BF Zhu , Y Liu . HLA-A, -B, and -DRB1 polymorphism defined by sequence-based typing of the Han population in Northern China . Tissue Antigens , 2006 . 67 ( 2 ): 146 - 152 . 16441486 DOI: 10.1111/j.1399-0039.2006.00529.x http://doi.org/10.1111/j.1399-0039.2006.00529.x . HLA-A, -B, and -DRB1 polymorphism defined by sequence-based typing of the Han population in Northern China ZL Yang , Y Guo , L Gao , Y Wang , YL Dong , WR Tang , KY Li , W Yan , CJ Xiao . Hypertension survey in Yi ethnic group in Yunnan Province, China . Zhonghua Liu Xing Bing Xue Za Zhi , 2004 . 25 ( 9 ): 817 (in Chinese) 15555370 . Hypertension survey in Yi ethnic group in Yunnan Province, China R Yuliwulandari , K Kashiwase , H Nakajima , J Uddin , TP Susmiarsih , AS Sofro , K Tokunaga . Polymorphisms of HLA genes in western Javanese (Indonesia): close affinities to Southeast Asian populations . Tissue Antigens , 2009 . 73 ( 1 ): 46 - 53 . 19140832 DOI: 10.1111/j.1399-0039.2008.01178.x http://doi.org/10.1111/j.1399-0039.2008.01178.x . Polymorphisms of HLA genes in western Javanese (Indonesia): close affinities to Southeast Asian populations H Zetterquist , O Olerup . Identification of the HLA-DRB1*04, -DRB1*07, and -DRB1*09 alleles by PCR amplification with sequence-specific primers (PCR-SSP) in 2 hours . Human Immunology , 1992 . 34 ( 1 ): 64 - 74 . 1356957 DOI: 10.1016/0198-8859(92)90086-3 http://doi.org/10.1016/0198-8859(92)90086-3 . Identification of the HLA-DRB1*04, -DRB1*07, and -DRB1*09 alleles by PCR amplification with sequence-specific primers (PCR-SSP) in 2 hours BF Zhu , CM Shen , QJ Wu , YJ Deng . Population data of 15 STR loci of Chinese Yi ethnic minority group . Legal Medicine (Tokyo) , 2008 . 10 ( 4 ): 220 - 224 . DOI: 10.1016/j.legalmed.2007.12.004 http://doi.org/10.1016/j.legalmed.2007.12.004 . Population data of 15 STR loci of Chinese Yi ethnic minority group 10.1631/jzus.B0900232 Project supported by the National Natural Science Foundation of China (Nos. 30700470 and 30871348), and the Shaan’xi Provincial Science and Technology Research and Development Project Fund (No. 2008K09-02), China 2009-08-01 2010-01-07 2009-11-18 2022-01-05 Copyright © Zhejiang University and Springer-Verlag Berlin Heidelberg 2010 2010 Journal of Zhejiang University-SCIENCE B (Biomedicine and Biotechnology) 02 11