5, -DQB1 alleles and their association with serologically defined HLA-A, -B, -C, -DR, and -DQ antigens

The HLA Dictionary 2001: A Summary of HLA-A, -B, -C, -DRB1/3/4/5, -DQB1 Alleles and Their Association With Serologically Defined HLA-A, -B, -C, -DR, and -DQ Antigens G. M. Th. Schreuder, C. K. Hurley, S. G. E. Marsh, M. Lau, M. Maiers, C. Kollman, and H. J. Noreen ABSTRACT: This report presents the serologic equivalents of 123 HLA-A, 272 HLA-B, and 155 HLA-DRB1 alleles. The equivalents cover over 64 percent of the presently identified HLA-A, -B, and -DRB1 alleles. The dictionary is an update of the one published in 1999 (Schreuder GMTh, Hurley CK, Marsh SGE, Lau M, Maiers M, Kollman C, Noreen H. The HLA dictionary 1999: a summary of HLA-A, -B, -C, -DRB1/3/4/5, -DQB1 alleles and their association with serologically defined HLA-A, -B, -C, -DR and -DQ antigens. Tissue Antigens 54:407, 1999) and also includes equivalents for HLA-C, DRB3, DRB4, DRB5, and DQB1 alleles. The data summarize information obtained by the WHO Nomenclature Committee for Factors of the HLA System, the International Cell Exchange (UCLA), the National Marrow Donor Program (NMDP), and individual laboratories. In addition, a listing is provided of alleles which are expressed as antigens with serologic reaction patterns that differ from the well-established HLA specificities.

INTRODUCTION Several years ago, the World Marrow Donor Association (WMDA) initiated a study aimed at identifying the serological types associated with each HLA-A, -B, -DRB allelic product. This work resulted in a “serology to DNA equivalents dictionary” first published in 1997

From the World Marrow Donor Association, Leiden, the Netherlands, Quality Assurance Working Group (G.M.Th.S., C.K.H., S.G.E.M., M.L.), WHO Nomenclature Committee for Factors of the HLA system (G.M.Th.S.., S.G.E.M.), International Cell Exchange, UCLA, Los Angeles, California (M.L.), US National Marrow Donor Program, Minneapolis, MN, USA, HLA Testing Laboratories (C.K.H., M.M., C.K., H.J.N.) and M. Setterholm. Address reprint requests to: G.M.Th. Schreuder, Ph.D., Department of Immunohematology and Blood Transfusion, E3-Q, Leiden University Medical Center, P.O. Box 9600, NL-2300 RC LEIDEN, The Netherlands; Tel: ⫹31-71-5263824; Fax: ⫹31-71-5248203; E-Mail: g.m.T. [email protected]. Received April 23, 2001; accepted April 24, 2001. Human Immunology 62, 826 – 849 (2001) © American Society for Histocompatibility and Immunogenetics, 2001 Published by Elsevier Science Inc.

The equivalents provided will be useful in guiding searches for unrelated hematopoietic stem cell donors in which patients and/or potential donors are typed by either serology or DNA-based methods. These equivalents will also serve typing and matching procedures for organ transplant programs where HLA typings from donors and from recipients on waiting lists represent mixtures of serologic and molecular typings. The tables with HLA equivalents and a questionnaire for submission of serologic reaction patterns for poorly identified allelic products will also be available on the WMDA web page: www.worldmarrow.org. Human Immunology 62, 826 – 849 (2001). © American Society for Histocompatibility and Immunogenetics, 2001. Published by Elsevier Science Inc. KEYWORDS: alleles; antigens; HLA; histocompatibility; serology

with an update in 1999 [1,2]. The dictionary is an attempt to aid searches for unrelated hematopoietic stem cell (hsc) donors in adult volunteer and umbilical cord blood banks. While most patients in need of hsc transplantation are HLA typed by DNA-based methods at medium or high resolution, substantial parts of the donor registries provide serologically based HLA typings only, at least for HLA class I. In this respect, the dictionary can help in the search for donors whose HLA phenotypes closely resemble that of the patient even though these typings are determined by different methods. Once identified, molecular class I typing of patient and selected donor can be performed to confirm the match. Since the appearance of the 1999 dictionary [2] based on the alleles listed in the 1998 WHO Nomenclature report [3], a large number of additional alleles have received official allele designations [4]. Moreover, a 0198-8859/01/$–see front matter S0198-8859(01)002471-3

HLA Alleles and Serologic Types

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TABLE 1 Comparison of all presently known HLA alleles as listed in the WHO nomenclature reports 1998 [3] and 2000 [4] and their known serologic equivalents as given in the previous [2] and the present “dictionary” 1999

Locus HLA-A HLA-B HLA-DRB1

2001

WHO Serologic WHO Serologic alleles [3] equivalents [2] alleles [4] equivalents N N (%) N N (%) 119 245 201

90 (76) 190 (77) 145 (72)

187 344 243

123 (65) 272 (79) 155 (64)

substantial number of DNA based typings have been added to the NMDP database prompting an update of the dictionary. As summarized in Table 1, serological equivalents are available for over 64% of the presently identified HLA-A, -B, and -DRB1 alleles. For completeness, all known alleles as published in the Nomenclature Report 2000 [4], including null alleles, have been included in Tables 2–7. Although serological typing of HLA-C and -DQ antigens is notoriously difficult, often incomplete and unreliable, the information, when available for a donor, may help in the selection procedure. For that reason we have included HLA-C and -DQB1 serologic equivalents. The amount of data for these loci is substantially increased by the inclusion of data from the NMDP. Although HLA-DQ molecules are heterodimers combining polymorphic alpha and beta chains, the DQ serologic patterns correlate strongly with DQB1, but not with DQA1 polymorphisms. For that reason, Table 7 gives equivalents between DQB1 alleles and DQ antigens. WHO NOMENCLATURE: ALLELE AND SEROLOGIC ASSIGNMENTS This update includes all presently identified alleles as included in the most recent nomenclature report [4]. Alleles are presented with their four-digit number designation, because silent substitutions (represented by the fifth digit) do not influence the antigenic expression. Nonexpressed (N-(ull))-alleles are included for completeness. The serologic assignment (second column of Tables 2–7) is taken from the Nomenclature Report 2000 as well. It should be noted that the latter report now incorporates data from the previous dictionaries for welldefined antigens only. In the Nomenclature Report 2000, serologic specificities of newly identified alleles are only included when reported by the allele contributors as being identical or very similar to the standard specificity associated with the allele group (e.g., A1 associated with

A*01 alleles). Likewise, the contributor may report an allele to be expressed as a serologically variant antigen. Such information is included in the nomenclature report under the heading “previous equivalents” and is included here in the comments column of Tables 2–7. Further information on the variant, if available, is included in Table 8. Some alleles have been assigned “standard” WHO HLA specificities although they were known to be slightly different. For example, the antigen encoded by B*1524 was assigned B62, but known to express the Bw4 instead of the Bw6 antigenic determinant present on the “standard” B62 [5]. INTERNATIONAL CELL EXCHANGE (UCLA) The HLA class I analysis included 88 samples, cells 977–1064, from healthy individuals, characterized between October 1998 and November 2000. Each cell was typed serologically by an average number of 201 laboratories, ranging from a low of 168 to a high of 229. The parallel typing for HLA-A, -B, -C alleles using DNAbased methods was performed by an average of 47 laboratories, ranging from 37 to 51 reporting monthly results. Some of the cells were reexamined in the DNA Extract-Class I Typing Exchange. Their high-resolution types were included in this update, together with the correlating serologically defined antigens of the initial typings. The number of participating labs typing the DNA extracts increased from 21 laboratories in 1997 to 60 laboratories in 2000. The average number was 54 laboratories during the time period used for this present update. The HLA class II analysis included 44 B lymphoblastoid cell lines (TER229-272) typed during the period between October 1998 and October 2000. The previously published listings [1,2] have been updated with these alleles (Tables 2–7). During this time period, each cell was typed serologically by an average number of 33 laboratories, ranging from a low of 27 to a high of 44. The parallel typing for HLA-DRB1, -DRB3, -DRB4, -DRB5, and DQB1-alleles using DNA-based methods was performed by an average of 99 laboratories ranging from a low of 92 to a high of 103. HLA serologic equivalents have been analyzed in heterozygous samples only if the other HLA allelic product would not interfere with the serologic recognition. For example, B*4010 expression was not analyzed on a cell also expressing B*4001 (cell 914⫽998). The associated serologic specificities as reported by the participating laboratories are given with the lowest and highest percentage recognition. The percentage recognition of the serological specificities has increased considerably. Therefore the percentage range as observed in the update analysis of 88 cells for

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TABLE 2 HLA-A alleles and their serologic designations International cell exchange UCLA

NMDP

HLA allele

WHO assigned

Cells tested

Assigned type (%)

Cells tested

Assigned type (%)

A*0101 A*0102 A*0103 A*0104N A*0105N A*0106 A*0107 A*0108 A*0201 A*0202 A*0203 A*0204 A*0205 A*0206 A*0207 A*0208 A*0209 A*0210 A*0211 A*0212 A*0213 A*0214 A*0215N A*0216 A*0217 A*0218 A*0219 A*0220 A*0221 A*0222 A*0224 A*0225 A*0226 A*0227 A*0228 A*0229 A*0230 A*0231 A*0232N A*0233 A*0234 A*0235 A*0236 A*0237 A*0238 A*0239 A*0240 A*0241 A*0242 A*0243N A*0244 A*0245 A*0246 A*0247 A*0248 A*0249 A*0301

A1 A1 A1 Null Null – A1 A1 A2 A2 A203 A2 A2 A2 A2 A2 A2 A210 A2 A2 A2 A2 Null A2 A2 A2 – A2 A2 A2 A2 A2 – – – A2 – A2 Null – A2 – – – – – – A2 A2 Null – – A2 – – – A3

14 – 1 – – – – – 52 – 6 – 2 16 4 – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – 22

A1[99–100] – A1 [99] – – – – – A2[99–100] – A2 [100] – A2 [100] A2 [100] A2 [100] – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – A3 [100]

4920 122 9 – – 1 – – 3527 735 361 16 301 276 71 2 2 2 191 3 2 7 – 18 33 1 – 3 – 12 3 – – – – – 1 – – – – – 1 – – – – – – – – – – – – – 2807

A1 A1 A1 – – A1 – – A2 A2 A2 A2 A2 A2 A2 A2 § A2 A2 A2 A2 A2 – A2 A2 A2 – A2 – A2 A2 – – – – – § – – – – – A2 – – – – – – – – – – – – – A3

[96] [92] [78]

Comments

A1[100]$

[100]@ [97] [81] [62] A28 [1] [69] [84] [67] [71] [50]@

A2 [94] A203 [6]$

[51]@ [73] [100]@ [51]@ [71] [50] [68] [100]@ [68]@ [75] [100]@

[100]@

(9)

[95]

HLA Alleles and Serologic Types

829

TABLE 2 (Continued) International cell exchange UCLA

NMDP

HLA allele

WHO assigned

Cells tested

Assigned type (%)

Cells tested

A*0302 A*0303N A*0304 A*0305 A*0306 A*0307 A*0308 A*1101 A*1102 A*1103 A*1104 A*1105 A*1106 A*1107 A*1108 A*1109 A*2301 A*2302 A*2303 A*2304 A*2305 A*2306 A*2402 A*2402102L A*2403 A*2404 A*2405 A*2406 A*2407 A*2408 A*2409N A*2410

A3 Null A3 A3 – – – A11 A11 A11 A11 A11 – A11 – – A23(9) – – – – – A24(9) Low A24 A2403 A24(9) A24(9) A24(9) A24(9) A24(9) Null A9

1 – – – – – – 30 1 – 2 – – – – – 17 – – – – – 47 – 7 – – – 11 – – –

A3 [100] – – – – – – A11 [99–100] A11 [99] – A11 [63–69] – – – – – A23 [98–100] – – – – – A24 [95–100] – A24 [44–67] A2403 [20–52] – – – A24 [99] – – –

287 – – 1 – – – 1757 23 13 3 2 – – – – 4807 2 – – – – 2209 – 239 10 10 3 232 33 – 29

A*2411N A*2413 A*2414 A*2415 A*2416 A*2417 A*2418 A*2419 A*2420 A*2421 A*2422 A*2423 A*2424 A*2425 A*2426 A*2427 A*2428 A*2501 A*2502 A*2503 A*2601 A*2602 A*2603 A*2604

Null A24(9) A24(9) – – – – – – – A9 A24(9) – – – A24(9) – A25(10) A10 – A26(10) A26(10) A26(10) A26(10)

– – – – – – – – – – – – – – – – – 5 – – 11 – 6 –

– – – – – – – – – – – – – – – – – A25 [94–99] – – A26 [90–98] – A26 [81–92] –

– 1 11 – – – – – 1 – 1 – – – – – – 1226 2 – 1497 20 31 1

Assigned type (%) A3 [76] – – blank @ – – – A11 [98] A11 [88] A11 [92] A11 [33] blank [67]@ A11 [100]@ – – – – A23 [96] A24 [1] A23 [100]@ – – – – A24 [96] A2403 [1] – A24 [50] blank [24] A9 [3] A2403 [3] A24 [60] A2403 [10] A24 [80] A23 [9] A24 [33] blank [33]@ A24 [61] blank [6] A24 [61] blank [21] – A24 [43] A9 [10] A2403 [3] blank [14] – § A24 [65] blank [18] – – – – – A24 [100]@ – A24 [100]@ – – – – – – A25 [95] A10 [2] A25 [100]@ – A26 [97] A10 [1] A26 [75] A34 [5] A26 [65] A10 [10] A26 [100]@

Comments

A11short A11var#

A24Neg$

A24var# A2403†

A31var# A24⫻3# A9short# A9var#

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TABLE 2 (Continued) International cell exchange UCLA HLA allele

WHO assigned

Cells tested

A*2605 A*2606 A*2607 A*2608 A*2609 A*2610 A*2611N A*2612 A*2613 A*2614 A*2615 A*2616 A*2617 A*2901 A*2902 A*2903 A*2904 A*3001 A*3002 A*3003 A*3004 A*3006 A*3007 A*3008 A*3009 A*3101 A*3102 A*3103 A*3104 A*3105 A*3201 A*3202 A*3203 A*3204 A*3205 A*3206 A*3301 A*3303 A*3304 A*3305 A*3401

A26(10) A26(10) A26(10) A26(10) A26(10) A10 Null – – – – – – A29(19) A29(19) – – A30(19) A30(19) A30(19) A30(19) – – – – A31(19) – – A31(19) A31(19) A32(19) A32(19) – – – – A33(19) A33(19) – – A34(10)

– – – – – – – – – – – – – – 5 – – 20 7 – – – – – – 21 – – – – 18 – – – – – 5 15 – – 9

A*3402 A*3403 A*3404

A34(10) – –

A*3601 A*4301 A*6601

Assigned type (%)

Cells tested

[87–95]

10 – – 49 2 – – – – – – – – 94 1212 3 – 2512 1770 6 50 – 2 – – 3209 – 1 – – 3136 3 1 3 – – 540 670 1 – 236

8 – –

A34 [93–95] – –

900 – –

A36 A43 A66(10)

3 – 8

769 8 712

A*6602

A66(10)

6

A*6603

A10

–

A36 [91–93] – A‘6601’ [45–57] A66 [17–39] A26 [12–19] A‘6602’ [53–65] A66 [15–18] A34 [7–15] –

A*6604 A*6801 A*6802 A*6803

– A68(28) A68(28) A68(28)

– 16 4 4

– – – – – – – – – – – – – – A29 – – A30 A30 – – – – – – A31 – – – – A32 – – – – – A33 A33 – – A34

NMDP

[98–99] [95–99] [96–99]

[97–99]

[97–99]

[95–97] [96–99]

– A68 [47–61] A28 [37–48] A68 [55–63] A28 [36–44] A68 [42–50] A28 [45–53]

180 25 – 2526 2012 125

Assigned type (%) A26 [90] A25 [10] – – A26 [98] A66 [2] A26 [100]@ – – – – – – – – A29 [94] A29 [98] A29 [100]@ – A30 [88] A31 [3] blank [6] A30 [94] A31 [2] A30 [67] blank [17] A30 [93] – A30 [100]@ – – A31 [90] A30 [3] A19 [1] blank [5] – A31[100]@ – – A32 [94] A32 [67]@ A32 [100]@ A3 [33] blank [33]@ – – A33 [87] A19 [3] A33 [94] A33 [100]@ – A34 [67] A26 [11] A10 [5] A66 [4] A33 [1] blank [8] A34 [90] A33 [3] A26 [2] A10 [2] – – A36 [68] A1 [16] blank [8] A26 [39] A29 [12] A26 [61] A66 [20] A34 [8] A10 [4] A25 [1] A11 [1] A66 [41] A34 [24] A10[11] A74 [7] A26 [3] A66 [36] A34 [16] A10 [9] A26 [8] A25 [4] A74 [4] – A28 [57] A68 [33] A28 [70] A68 [21] A69 [1] A28 [62] A68 [28]

Comments

A10short# A10var#

A3#

A34 [42] 34⫻31 [25] blank [33]$#

A10short#

HLA Alleles and Serologic Types

831

TABLE 2 (Continued) International cell exchange UCLA HLA allele

WHO assigned

Cells tested

A*6804 A*6805 A*6806 A*6807 A*6808 A*6809 A*6810 A*6811N A*6812 A*6813 A*6814 A*6815 A*6816 A*6817 A*6818N A*6819 A*6901 A*7401

A68(28) A68(28) – – A68(28) – – Null A28 – – – A68(28) – Null – A69(28) A74(19)

– – – – 2 – – – – – – – – – – – 1 5

– – – – A68 [38–45] A28 [53] – – – – – – – – – – – A28 [49] A69 [40] A2 [8] A74 [77–85]

A*7402 A*7403

A74(19) A74(19)

– 3

– A74 [86–89]

A*7404 A*7405 A*8001

– – A80

– – 7

– – A80 [69–81] A36 [4–5]

Assigned type (%)

NMDP Cells tested 6 46 2 – – – 2 – 2 – – – – – – – 252 88 2 28 – – 480

Assigned type (%) A28 [52] A68 [17] A28 [49] A68 [33] A28 [50] A68 [50]@ – – – § – § – – – – – – – A28 [53] A69 [9] A2 [3] A68 [3] A74 [42] blank [28] A19 [8] A33 [3] A32 [2] A31 [2] A74 [50] A33 [50]@ blank [67] A74 [11] A19 [7] A29 [4] A33 [4] – – blank [70] A80 [10] A’X’ [8] A36 [2] A1 [1]

Comments

A28short#

@ ⫽ allele has been reported ⬍6 times and/or serologically identified in ⬍4 individuals. § ⫽ allele identified but serology not informative. # ⫽ see remarks in Table 8. $ ⫽ HLA-Club Cell Exchange: one sample typed by 17 laboratories and allele identified by at least 3 laboratories. † ⫽ locally identified in Leiden.

Class I and 44 cells for Class II are now presented in the tables. The International Cell Exchange (UCLA) provided up to now serologic equivalents for 39 HLA-A, 99 HLA-B, 29 HLA-Cw, 58 HLA-DRB1, 6 HLA-DRB3, 3 HLA-DRB4, 3 HLA-DRB5, and 17 HLA-DQB1 alleles. NATIONAL MARROW DONOR PROGRAM (NMDP) Cells from bone marrow transplant patients and unrelated donors were HLA typed using serological and molecular methods by typing laboratories participating in a project to retrospectively type transplant pairs [6] and by laboratories associated with NMDP-affiliated donor or transplant centres as outlined in the previous report [2]. The NMDP dataset is based on the typing information from 35,102 individuals for HLA-A; 56,718 individuals for HLA-B; and 65,752 individuals HLA-DRB1. Anal-

ysis was extended to HLA-C (4,749 individuals) and HLA-DQB1 (36,297 individuals). Tables 2–5 and 7 present the number of individuals carrying a certain allele, together with the most frequent serologic type(s) associated with that allele. For example (Table 2), there were 4,920 individuals typed as A*0101 and most (96%) received the HLA-A1 serologic assignment. Other alleles appear with several common assignments. For example, B*5108 was observed in 131 individuals of which 77% were serologically typed as B51, 4% as B53, 2% as B5, and 2% as B52. The EM algorithm used to obtain these data has been described previously [2]. If an allele appeared five times or less, the serologic assignment(s) should be considered as only an approximation of the serologic reactivity of the resultant antigen. These cases are indicated in the tables. The NMDP data presented in the tables lacks information on the actual serologic reaction patterns which might vary from the standard assignment.

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TABLE 3 HLA-B alleles and their serologic designations International cell exchange UCLA HLA allele

WHO Cells assigned tested

B*0702 B*0703 B*0704 B*0705 B*0706 B*0707 B*0708 B*0709 B*0710 B*0711 B*0712 B*0713 B*0714 B*0715 B*0716 B*0717 B*0718 B*0719 B*0720 B*0721 B*0722 B*0723 B*0724 B*0725 B*0726 B*0801 B*0802 B*0803 B*0804 B*0805 B*0806 B*0807 B*0808N B*0809 B*0810 B*0811 B*0812 B*0813 B*1301 B*1302 B*1303 B*1304

B7 B703 B7 B7 B7 B7 – B7 – B7 – – – B7 B7 – – – – – – – B7 – – B8 B8 B8 – – B8 B8 Null B8 B8 – – – B13 B13 – –

20 – – 2 – – – 2 – – – – – – – – – – – – – – – – – 1 – – 3 – – – – – – – – – 2 3 – 2

B*1306 B*1307 B*1401

– Null B64(14)

– – 3

B*1402 B*1403 B*1404 B*1405 B*1406 B*1501 B*1502

B65(14) B14 – – B14 B62(15) B75(15)

3 – – – – 13 12

B*1503

B72(70)

12

B*1504 B*1505

B62(15) B62(15)

– 1

Assigned type (%) B7 [96–99] – – B7 [75–99] – – – B7 [97–98] – – – – – – – – – – – – – – – – – B8 [99] – – B8 [51–71] B59 [6–7] – – – – – – – – – B13 [100] B13 [99–100] – B21 [22–38] B49 [15] B15 [7–17] B15⫻21[0–15] B50 [5–11] B63 [0–8] B77 [4–7] – – B14 [49–50] B64 [26–37] B65 [11–22] B65 [65–68] B14 [30–34] – – – – B62 [92–98] B75 [83–88] B15 [5–10] B62 [6– 7] B70 [50–59] B72 [28–37] – B62 [88]

NMDP Cells tested 10283 14 42 59 8 12 – 7 – – 5 3 1 1 – – – – – – – – – – – 6351 2 – 6 1 1 – – 1 – – – – 210 944 1 –

Assigned type (%) B7 [98] B7 [93] B7 [89] B7 [94] B7 [100] B7 [81] – B7 [83] – – B7 [100]@ blank [100]@ B7 [100]@ B7 [100]@ – – – – – – – – – – – B8 [97] § – blank [25] B8 [17] § § – – B8 [100]@ – – – – B13 [97] B13 [99] B13 [100]@ –

Comments B703#

B7short#

B7var# B7short#

B7short#

B7weak#

B8var# B8var# B8var# (10) (7,10) B8var#

B21var# B15⫻21# B’X’ [53] B15[12] B21 [12]$

– – – – 649 B14 [69] B65 [14] B64 [9] 1542 37 2 2 – 1496 883

B14 [63] B65 B14 [49] B65 B14 [50] B64 blank [100]@ – B62 [94] B15 B75 [65] B62

[33] B64 [1] [35] B64 [3] [50]@ B14weak(4) [3] [20] B15 [6]

2295 B70 [60] B72 [10] B35 [2] B62 [1] blank [17] 21 B62 [71] B75 [10] B15 [2] blank [11] 19 B62 [58] B15 [21] B75 [11]

HLA Alleles and Serologic Types

833

TABLE 3 (Continued) International cell exchange UCLA HLA allele

WHO Cells assigned tested

Assigned type (%)

B*1506 B*1507 B*1508

B62(15) B62(15) B75(15)

– 1 2

– B62 [93] B75 [30–48] B15 [19–35] B62 [6–19] B‘1508’ [11–14] – B70 [60–67] B71 [22–33]

B*1509 B*1510

B70 B71(70)

– 5

B*1511

B75(15)

3

B*1512

B76(15)

5

B75 [31–51] B15 [21–38] B46 [6–12] B‘1511’ [0–19] B76 [76–88] B45 [3–6]

B*1513

B77(15)

2

B77 [46–53] B63 [0–16]

B*1514

B76(15)

1

B*1515

B62(15)

4

B*1516 B*1517 B*1518 B*1519

B63(15) B63(15) B71(70) B76(15)

7 3 7 1

B*1520 B*1521

B62(15) B75(15)

– 6

B*1522 B*1523

B35 –

1 2

B*1524

B62(15)

4

B*1525 B*1526N B*1527 B*1528 B*1529

B62(15) Null B62(15) B15 B15

5 – – – 2

B*1530

B62(15)

3

B*1531 B*1532 B*1533 B*1534 B*1535 B*1536 B*1537

B75(15) B62(15) B15 B15 B62(15) – B70

– – – – 5 – 2

B76 [23] B45 [20] B15 [20] B70 [6] B63 [5] B75 [51–59] B62 [15–29] B15 [13–27] B63 [87–91] B63 [89–95] B70 [57–69] B71 [19–31] B76 [29] B15 [19] B45 [11] B62 [8] B75 [7] – B75 [78–86] B15 [12–13] B62 [4–8] B35 [79] B70 [12] B15 [5] B‘1523’ [17] B5 [14–15] B77 [7–11] B70 [7–9] B15 [5–8] B62 [45–48] B‘1524’ [23–24] B15 [21–22] B77 [4–6] B63 [2–3] B62 [86–95] – – – B70 [54–58] B15 [11–12] B71 [7–10] B62 [63–86] B75 [4–13] B15 [12–19] – – – – B62 [88–95] – B70 [46] B71 [11] B78 [9]

B*1538 B*1539 B*1540 B*1542 B*1543 B*1544 B*1545 B*1546 B*1547 B*1548 B*1549 B*1550 B*1551 B*1552

– B62(15) – – – – B62(15) B72(70) – B62(15) – – B70 –

– – – – – – – – – – – – – –

– – – – – – – – – – – – – –

NMDP Cells tested

Assigned type (%)

Comments

10 B62 [40] B70 [20] B75 [11] 85 B62 [78] B15 [1] B75 [1] 42 B62 [21] B15 [17] B75 [17] B35 [10] B70 [7] 63 B70 [51] B35 [3] B15 [2] blank [20] 1051 B70 [46] B35 [4] B72 [2] B71 [2] B75 [1] B15 [1] blank [18] 43 B75 [26] B62 [25] B46 [19] B15 [14] blank [9] 9 B76 [22] B75 [11] B62 [11] B15 [11] B53 [11] 91 B63 [24] B77 [19] B75 [15] B15 [13] B’X’ [11] B62 [8] B76 [1] 3 B62 [33]@ 95 B62 [48] B75 [33] B15 [11] B70 [4] 706 651 491 –

B75 (11)

B63 [81] B57 [4] B15 [3] B62 [1] B’X’ [2] B63 [88] B15 [3] B57 [1] B70 [49] blank [39] B71 [4] B72 [1] –

4 B62 [100]@ 119 B75 [56] B62 [8] B15 [7] B35 [5] B’X’ [5] B70 [3] 311 B35 [61] B70 [13] B15 [2] B53 [1] B78 [1] 11 blank [64] B70 [18] B53 [9] 51 B62 [51] B’X’ [7] B77 [4] B63 [2] B15 [2] blank [16] 84 – 71 6 20

B77 [59] B62 [29]$

B75[76]$ NM5, Bw4pos# B62–Bw4#

B62 [76] B15 [9] B75 [6] – B62 [75] B15 [7] B62 [67] B15 [17] B75 [17] blank [60] B70 [20] B35 [6]

82 B62 [63] B15 [20] B75 [6] 8 – – 1 17 3 23 4 16 22 – – – 16 4 15 – – – – –

B62 [63] B70 [13] – – B62 [100]@ B62 [86] B15 [6] B13 [100]@ blank [44] B70 [18] B’X’ [17] B15 [4] B71 [4] B78 [4] B62 [75]@ B62 [68] B15 [13] blank [67] B70 [12] B62 [5] – – – B62 [46] B50 [50] blank [53] B35 [13] B70 [9] B15 [7] – – – – –

Bw4pos B15short# B62var# B62short (11) Bw4pos

(10)

834

G. M. Th. Schreuder et al.

TABLE 3 (Continued) International cell exchange UCLA HLA allele

WHO assigned

Cells tested

B*1553 B*1554 B*1555 B*1556 B*1557 B*1558 B*1559 B*1560 B*1561 B*1562 B*1563 B*1564 B*1801 B*1802 B*1803 B*1804 B*1805 B*1806 B*1807 B*1808 B*1809 B*1810 B*1811 B*1812 B*1813 B*2701 B*2702 B*2703 B*2704 B*2705 B*2706 B*2707 B*2708

– – B15 – – B62(15) B35 – – – – – B18 B18 B18 – B18 B18 – – B18 – – – – B27 B27 B27 B27 B27 B27 B27 B2708

– – – – – – – – – – – – 7 – – – – – – – – – – – – – 1 1 – 3 2 – 3

B*2709 B*2710 B*2711 B*2712

B27 B27 B27 –

– – – –

B*2713 B*2714 B*2715 B*2716 B*2717 B*2718 B*2719 B*2720 B*2721 B*2722 B*2723 B*3501 B*3502 B*3503 B*3504 B*3505 B*3506 B*3507 B*3508 B*3509

B27 – – – B27 – B27 B27 – B27 – B35 B35 B35 B35 B35 B35 B35 B35 B35

– – – – – – – – – 1 – 13 2 2 – 7 – – 4 –

Assigned type (%)

NMDP Cells tested

– – – – – – – – – – – – B18 [97–98] – – – – – – – – – – – – – B27 [97] B27 [97] – B27 [98–100] B27 [99–100] – B2708 [36–58] B7 [19–41] B27 [11–19] – – – –

– – – – – – – – – – – – 1431 27 27 2 – – 1 1 – – – – – 6 351 19 15 1100 21 6 13

– – – – – – – – – B27 – B35 B35 B35 – B35 – – B35 –

– 4 – – – – – – – – – 1252 766 904 15 154 9 – 217 –

[100] [94–99] [94–97] [95–100] [96–99] [95–99]

4 1 2 14

Assigned type (%) – – – – – – – – – – – – B18 B18 B18 B18 – – § B18 – – – – – B27 B27 B27 B27 B27 B27 B27 B27

B15–Bw6, Ref. [12] Ref. [9] B35, Ref. [13]

[96] [89] [85] blank [15] [100]@ B18var# [100]@ B18–Bw4#

[100] [85] blank [7] [95] [100] [98] [95] [100] [46] B7 [15] blank [31]

B27 [100]@ B27 [100]@ B27 [100]@ blank [86] – B27 – – – – – – – – – B35 B35 B35 B35 B35 B35 – B35 –

Comments

B27[100]$, B7Qui#

B27var# BX [63] B40 [25] B61 [12]$#

[75]@ B’X’#

Ref. [9] Ref. [14] [96] [68] [96] [93] [97] [77] [94]

HLA Alleles and Serologic Types

835

TABLE 3 (Continued) International cell exchange UCLA HLA allele

WHO Cells assigned tested

B*3510 B*3511 B*3512 B*3513 B*3514 B*3515 B*3516 B*3517 B*3518 B*3519 B*3520 B*3521 B*3522 B*3523 B*3524 B*3525 B*3526 B*3527 B*3528 B*3529 B*3530 B*3531 B*3532 B*3533 B*3534 B*3535 B*3536 B*3537 B*3701 B*3702 B*3703N B*3801 B*3802 B*3803 B*3804 B*3805 B*3806 B*3807 B*3901 B*3902 B*3903 B*3904 B*3905

B35 B35 B35 B35 B35 B35 B35 B35 B35 B35 B35 – – – – – – B35 – B35 B35 – B35 – – B35 – – B37 – Null B38(16) B38(16) B16 – B38(16) – – B3901 B3902 B39(16) B39(16) B16

– – 2 – – – 2 3 – – – – – – – – – – – – – – – – – – – – 3 – – 3 3 – – – – – 4 2 – – 6

B*3906 B*3907 B*3908 B*3909 B*3910 B*3911 B*3912 B*3913 B*3914 B*3915 B*3916 B*3917 B*3918 B*3919 B*3920

B39(16) B39(16) B39(16) B39(16) B39(16) B39(16) B39(16) B39(16) – – – – – – –

3 – 1 – 2 – – – – – – – – – –

Assigned type (%) – – B35 [72–80] B53 [11–16] – – – B35 [63] B70 [20] B35 [98–99] – – – – – – – – – – – – – – – – – – – – B37 [98–99] – – B38 [97–98] B38 [90–91] – – – – – B39 [99] B39 [90–97] B67 [0–5] – – B39 [69–90] B16 [3–11] B38 [3–9] B‘3905’ [3–8] B39 [99] – B39 [48] B16 [12] – B39 [95–99] – – – – – – – – – –

NMDP Cells tested

Assigned type (%)

10 17 347 – 15 5 33 195 1 3 16 7 1 3 5 1 – 1 4 1 – 3 – – – – – – 1622 4 – 1874 641 – – – – – 308 71 16 1 150

B35 B35 B35 – B35 B35 B35 B35 B35 B35 B35 B35 B35 B35 B35 § – B35 B35 B35 – § – – – – – – B37 B27 – B38 B38 – – – – – B39 B39 B39 B39 B39

371 3 48 4 106 13 – – 2 3 – – – – –

B39 B39 B39 B39 B39 B39 – – B39 B39 – – – –

[80] [67] B51 [10] [78] [87] [40] blank [60] [37] B’X’ [15] B70 [9] blank [27] [95] [100]@ [33] B21 [33] blank [33]@ [39] B70 [6] blank [12] [22] B78 [14] blank [43] [100]@ [34] blank [66]@ [23] B78 [20]@ [100]@ [50] blank [50]@ [100]@

Comments B35short, Ref. [11]

B35var#

(10) Bfu/B40–like# (10)

[93] Blank [6] [50] B37 [25] B’X’ [25]@

B’blank’#

[93] B16 [1] B39 [1] [90] B39 [2] B16 [2] blank [6] #

[91] B3901 [6] B38 [1] B16 [1] [85] blank [11] [78] [100]@ [76] B38 [3] B16 [1] blank [7] [93] [36]@ [46] B’X’ [12] blank [29] [79]@ [84] B16 [1] B38 [1] blank [14] [62] blank [23] [100]@ [100]@

B39var#

B39, Ref. [11] B39, Ref. [11]

836

G. M. Th. Schreuder et al.

TABLE 3 (Continued) International cell exchange UCLA HLA allele

WHO Cells assigned tested

B*3922 B*3923 B*3924 B*4001 B*4002 B*4003 B*4004 B*4005

– B39(16) B39(16) B60(40) B61(40) B61(40) B61(40) B4005

– – – 23 9 – – 3

B*4006 B*4007 B*4008

B61(40) B60(40) –

2 – 2

B*4009 B*4010 B*4011 B*4012 B*4013 B*4014 B*4015 B*4016 B*4018 B*4019 B*4020 B*4021 B*4022N B*4023 B*4024 B*4025 B*4026 B*4027 B*4028 B*4029 B*4030 B*4031 B*4032 B*4033 B*4034 B*4035 B*4101 B*4102 B*4103 B*4104 B*4105 B*4201 B*4202 B*4402 B*4403 B*4404 B*4405 B*4406 B*4407 B*4408 B*4409 B*4410 B*4411 B*4412 B*4413

B61(40) B60(40) B40 – – – – B61 (40) – – – – Null – – – B21 B61(40) – B61(40) – B60(40) – – B60(40) – B41 B41 B41 – – B42 B42 B44(12) B44(12) B44(12) B44(12) B44(12) B44(12) B44(12) B12 B44(12) – B44(12) B44(12)

– 1 – 2 – – – – – – – – – – – – – – – – – – – – – – 4 3 – – – 8 5 6 13 – – – – – – – – – –

Assigned type (%) – – – B60 [96–99] B61 [84–87] – – B4005 [41–57] B50 [28–36] B21 [5–9] B61 [81–86] B60 [5–7] – B61 [24–27] B40 [20–30] B48 [11–14] B‘4008’ [9] – B60 [60] B40 [19] B61 [12] B48 [7] – B48 [34] B70 [20] B72 [10] – – – – – – – – – – – – – – – – – – – – – – B41 [91–95] B41 [95–97] – – – B42 [92–96] B42 [87–96] B67 [0–12] B44 [99–100] B44 [99–100] – – – – – – – – – –

NMDP Cells tested – – – 3285 1056 23 43 257 413 12 127 4 2 8 8 1 2 – 3 2 – 4 – – – – – – 1 – – – – – – – – 479 390 10 – – 926 65 6374 2058 30 101 8 2 1 6 32 – – –

Assigned type (%)

Comments

– – – [15] B60 [93] B40 [3] B61 [2] B61 [80] B60 [9] B40 [6] B61 [62] B60 [22] blank [8] B61 [35] B50 [16] B60 [12] B40 [5] B50 [30] B4005 [13] blank [23] B’X’ [4] B61 BN21, B21var [7] B70 [5] B21 [5] B60 [5] B61 [74] B60 [5] B40 [5] blank [8] B60 [83] B40 [8] B’Fu’ blank [69] B61 [12] B40 [8] B48 [2] B40x48var# B61 [75]@ B60 [100]@ B61 [41] B60 [14] B40 [12] B60 [50] B48 [13] blank [38] § B60 [50] B61 [50]@ – B61 [67] B60 [33]@ B61 [100]@ – B61 [50] B60 [27]@ – – – – – – B61 [100]@ – – – – – – – – B41 [86] blank [11] B41 [87] blank [9] B41 [90] – – B42 [93] B55 [1] B7 [1] B42 [95] B44 [97] B44 [96] B44 [58] B12 [3] B45 [3] B44 [95] blank [50] B44 [37] B44 [100]@ B44 [100]@ B45 [83] B44 [78] B12 [3] B45 [3] – – –

B60var# B48⫻70# B40Bw4

Bw4pos B40⫻15#

B40short#

B44[100]$ B12var# B44var# B12Bw6#

HLA Alleles and Serologic Types

837

TABLE 3 (Continued) International cell exchange UCLA HLA allele

WHO assigned

Cells tested

B*4414 B*4415 B*4416 B*4417 B*4418 B*4419N B*4420 B*4421 B*4422 B*4423N B*4424 B*4501 B*4502 B*4503 B*4504 B*4601 B*4602 B*4701 B*4702 B*4703

– B12 B47 – – Null – – – Null – B45(12) – – – B46 B46 B47 B47 –

– – 1 – – – – – – – – 4 – – – 6 – 2 – 2

B*4801 B*4802 B*4803 B*4804 B*4805 B*4806 B*4807 B*4901 B*4902 B*4903 B*5001 B*5002 B*5004 B*5101 B*5102

B48 B48 B48 – B48 – B48 B49(21) – – B50(21) B45(12) B50(21) B51(5) B5102

6 – – – – – – 2 – – 7 – – 22 6

B*5103 B*5104 B*5105 B*5106

B5103 B51(5) B51(5) B5

– – – 1

B*5107 B*5108 B*5109 B*5110 B*5111 B*5112 B*5113 B*5114 B*5115 B*5116 B*5117 B*5118 B*5119 B*5120 B*5121 B*5122 B*5123

B51(5) B51(5) B51(5) – Null – – – – B52(5) B51(5) B51(5) – – – – –

– – – – – – – – – – – – – – – – –

Assigned type (%) – – B47 [52] B44 [12] – – – – – – – – B45 [96–99] – – – B46 [85–97] – B47 [90–94] – B47 [70–77] B60 [4–9] B61 [7–8] B40 [6–7] B48 [92–96] – – – – – – B49 [92–95] – – B50 [87–99] – – B51 [92–98] B51 [33–44] B5102 [28–31] B53 [14–21] B52 [2–5] – – – B51 [43] B53 [32] B78 [11] B5102 [7] B5 [7] – – – – – – – – – – – – – – – – –

NMDP Cells tested

Assigned type (%)

– – – – – – – – – – – 2073 3 – – 1410 – 295 6 3

– – – – – – – – – – – B45 B45 – – B46 – B47 B47 B47

490 39 23 3 – – 1 2550 – – 1527 71 – 1425 183

B48 [80] B60 [6] B61 [2] B40 [2] blank [5] blank [44] B35 [16] B70 [13] B48 [5] B48 [57] B40 [18] blank [13] B48 [67]@ – – B48 [100]@ B49 [94] B50 [1] B21 [1] – – B50 [84] B21 [3] B49 [2] B45 [2] B45 [92] B50 [3] – B51 [96] B5 [2] B52 [1] B51 [74] B53 [10] B52 [6] B5 [3] B5102 [3]

– 4 24 34 57 131 33 3 – – 3 4 – – – – – – – – –

B12Bw4# # B44 [17]

[94] B44 [1] B12 [1] B50 [1] [67]@ [59] blank [35] B’X’ [3] B62 [1] B46var# [52] B27 [4] blank [34] [33] B40 [34] blank [34] [33] B13 [33]@

– B51 [75] B5 [25]@ B51 [54] B53 [21] B52 [8] B51 [62] B53 [15] B52 [9] B5 [3] B51 B51 B51 § – – B51 B51 – – – – – – – – –

Comments

B47–Bw6# B47var#

B5var#

[80] B52 [9] [77] B53 [4] B5 [2] B52 [2] [88] B5 [3] B52 [3] B53 [3] B78 [3] B’blank’# [100]@ [100]@

838

G. M. Th. Schreuder et al.

TABLE 3 (Continued) International cell exchange UCLA HLA allele

WHO Cells assigned tested

Assigned type (%)

NMDP Cells tested

B*5124 B*5201 B*5202 B*5301 B*5302 B*5303 B*5304 B*5305 B*5306 B*5307 B*5401

B51(5) B52(5) – B53 – – – – – – B54(22)

– 10 – 11 – – – – – – 4

– B52 [90–96] – B53 [85–95] – – – – – – B54 [85–88]

– 2383 – 2749 3 1 – – – – 590

B*5402 B*5501 B*5502 B*5503 B*5504 B*5505 B*5507 B*5508 B*5509 B*5510 B*5601 B*5602 B*5603 B*5604 B*5605 B*5606 B*5607 B*5701 B*5702 B*5703 B*5704 B*5705 B*5706 B*5707 B*5801 B*5802 B*5804 B*5805 B*5806 B*5901 B*6701

B54(22) B55(22) B55(22) – B55(22) B22 B54(22) – – B55(22) B56(22) B56(22) B22 B56(22) B56(22) B78 B56(22) B57(17) B57(17) B57(17) B57(17) – – – B58(17) B58(17) – – – B59 B67

– 3 – – – – – – – – 3 – – 1 – – – 6 3 2 – – – – 13 1 – – – 4 2

– 314 249 1 3 – 2 – – – 581 5 13 8 3 – – 3335 52 289 30 2 – – 1767 766 – – – 118 76

B*7301 B73 B*7801 B78

3 4

– B55 [89–97] – – – – – – – – B56 [94–96] – – B56 [78] B22 [6] – – – B57 [97–98] B57 [96–99] B57 [90–94] – – – – B58 [85–92] B58 [82] – – – B59 [95–96] B67 [74] B39 [12] B42 [2–9] B55 [0–8] B73 [81–83] B78 [83–89] B35 [1–9] B51 [1–6]

B*7802 B*7803 B*7804 B*7805 B*8101 B*8201

– – – – 9 1

– – – – B81 [79–89] B7 [13–19] B‘8201’ [28] B22 [9]

– –

– –

B78 – – – B81 –

B*8202 – B*8301 –

99 182 – – 2 – 590 140 – –

Assigned type (%) – B52 [82] B51 [5] B5 [2] – B53 [87] B35 [5] B53 [67] B51 [33]@ B35 [100]@ – – – – B54 [68] blank [12] B55 [10] B56 [3] B22 [3] B’X’ [2] – B55 [97] B22 [2] B56 [1] B55 [84] B54 [3] B22 [3] B22 [100]@ B55 [67]@ – B54 [50]@ – – – B56 [73] B55 [3] B22 [3] blank [16] B56 [80]@ B56 [46] B22 [8] B56 [63] B7 [67]@ – – B57 [93] B17 [4] B58 [2] B57 [75] B58 [10] B17 [8] B57 [83] B17 [9] B58 [6] B57 [69] B17 [8] B57 [100]@ – – B58 [80] B57 [7] B17 [4] B58 [72] B57 [12] B17 [3] – – – B59 [44] blank [41] B38 [3] B55 [3] B39 [41] B67 [31] B22 [11] blank [11] blank [78] B73 [20] B78 [43] B35 [13] B5 [14] B’X’ [7] B70 [3] blank [15] – – B35 [51]@ – B7 [76] B81 [8] blank [6] B60 [1] B48 [1] blank [59] B’X’ [10] B82 [4] B22 [6] B70 [4] B45 [4] – –

@ ⫽ allele has been reported ⬍6 times and/or serologically identified in ⬍4 individuals. § ⫽ allele identified but serology not informative. # ⫽ see remarks in Table 8. $ ⫽ HLA–Club Cell Exchange: one sample typed by 17 laboratories and allele identified by at least three laboratories.

Comments (7)

B52/53# B51/53# B53⫻37#

B55/67#

B22short#

B22var# B56var# B78var# B56Bw4#

B22⫻45, B45var# B‘8201’

HLA Alleles and Serologic Types

839

TABLE 4 HLA–C alleles and their serologic designations International cell exchange ULCA HLA alleles

WHO assigned

Cells tested

Cw*0102 Cw*0103 Cw*0104 Cw*0202 Cw*0203 Cw*0204 Cw*0302 Cw*0303 Cw*0304 Cw*0305 Cw*0306 Cw*0307 Cw*0308 Cw*0309 Cw*0310 Cw*0311 Cw*0312 Cw*0401 Cw*0403

Cw1 Cw1 – Cw2 – – Cw10(w3) Cw9(w3) Cw10(w3) – – Cw3 – Cw3 Cw3 – – Cw4 –

33 – – 17 – – 5 23 28 – – – – 2 – – – 45 8

Cw*0404 Cw*0405 Cw*0406 Cw*0407 Cw*0408 Cw*0501 Cw*0502 Cw*0503 Cw*0504 Cw*0602 Cw*0603 Cw*0604 Cw*0605 Cw*0606 Cw*0607 Cw*0701 Cw*0702 Cw*0703 Cw*0704 Cw*0705 Cw*0706 Cw*0707 Cw*0708 Cw*0709 Cw*0710 Cw*0711 Cw*0712 Cw*0713 Cw*0714 Cw*0801 Cw*0802 Cw*0803 Cw*0804 Cw*0805 Cw*0806 Cw*0807 Cw*0808 Cw*0809

– – – – – Cw5 Cw5 – – Cw6 – – Cw6 – – Cw7 Cw7 – Cw7 – Cw7 – – – – – – – Cw7 Cw8 Cw8 Cw8 Cw8 – – – – –

– – 2 – – 7 – – – 18 – – – – – 17 42 – 11 – 3 – – – – – – – – 25 9 1 3 – – – – –

Assigned type (%) Cw1 [56–85] – – Cw2 [58–87] – – Cw3 [56–75] Cw3 [64–74] Cw3 [64–71] – – – – Cw3 [44–51] – – – Cw4 [58–82] Cw6 [18–40] C‘403’ [2–5] – – Cw6 [56–62] – – Cw5 [58–72] – – – Cw6 [52–74] – – – – – Cw7 [51–83] Cw7 [55–78] – Cw7 [62–80] – Cw7 [68–76] – – – – – – – – Cw8 [48–64] Cw8 [35–65] Cw8 [8] Cw8 [43–59] – – – – –

Cw10 [5–9] Cw9 [5–10] Cw10 [5–12]

Cw10 [3–5]

Cw6 [0–6] Cw4 [13–25]

Cw4 [8–13]

NMDP Cells tested

Assigned type (%)

292 1 – 442 – – 40 556 798 1 1 – – – 1 – – 998 3

Cw1 Cw1 – Cw2 – – Cw3 Cw3 Cw3 Cw3 Cw3 – – – Cw3 – – Cw4 Cw6

2 1 – – – 869 – – – 855 1 1 – – – 1227 1234 1 104 – 3 1 – – 1 – – – – 31 270 1 2 – – – – –

§ Cw4 – – – Cw5 – – – Cw6 Cw6 Cw6 – – – Cw7 Cw7 Cw7 Cw7 – Cw7 § – – Cw7 – – – – Cw8 Cw8 § Cw8 – – – – –

Comments

[92] [100]@ [91] [86] Cw10 [5] [86] Cw9 [4] [86] Cw10 [3] [100]@ [100]@

[100]@ [91] [67]@ [100]@

[82] blank [12]

[76] blank [18] [100]@ [100]@

[69] blank [14] [74] blank [12] [100]@ [65] blank [21] [80]@

[100]@

[45] blank [45] [65] blank [27] [50]@

Cw4NM, Cw4/6var

840

G. M. Th. Schreuder et al.

TABLE 4 (Continued) International cell exchange ULCA

NMDP

WHO assigned

Cells tested

Cw*1202 Cw*1203 Cw*1204 Cw*1205 Cw*1206 Cw*1207 Cw*1301 Cw*1402 Cw*1403 Cw*1404 Cw*1502

– – – – – – – – – – –

7 12 – – – – – 11 1 – 12

blank& Cw7 [2–14] – – – – – blank& blank& – Cw6 [0–11] Cw2 [0–7]

82 322 2 – – – – 102 8 – 131

Cw*1503 Cw*1504 Cw*1505 Cw*1506 Cw*1507 Cw*1508 Cw*1509 Cw*1510 Cw*1601 Cw*1602 Cw*1604 Cw*1701

– – – – – – – – – – – –

– – 6 – – – – – 22 – – 17

– – Cw2 [0–39] – – – – – Cw7 [0–13] – – C‘17’ [33–34] Cw7 [16–24]

1 – 28 1 – – 1 – 286 24 5 68

Cw*1702 Cw*1703 Cw*1801 Cw*1802

– – – –

– – 2 2

HLA alleles

Assigned type (%)

Cells tested

– – Cw6 [37–42] Cw6 [38–43]

– – 8 –

Assigned type (%)

Comments

blank [91] Cw7 [5] blank [61] § – – – – blank [84] Cw1 [4] blank [88] – blank [78] Cw2 [6] Cw6 [3] § – blank [29] Cw7 [23] § – – § – blank [92] blank [81] blank [78] @ Cw7 [41] Cw2 [11] blank [35] – – Cw6 [62] –

& ⫽ no Cw antigen reported by more than 5% of the laboratories. @ ⫽ allele has been reported ⬍6 times and/or serologically identified in ⬍4 individuals. § allele identified but serology not informative.

INFORMATION OBTAINED FROM OTHER SOURCES The comments column in the tables indicate references to recent publications that included remarks on the serologic expression of newly described alleles. Expression of alleles tested in the HLA-Club Cell Exchange are indicated in the tables as well. Also, personal communications to and local observations in the Leiden laboratory are included, but it should be realized that such observations are often based on very few samples tested in only one laboratory. SEROLOGIC SPECIFICITIES LACKING OFFICIAL WHO NOMENCLATURE DESIGNATIONS As shown in the tables, a number of alleles appear with several common serologic assignments, indi-

cating that their serologic reaction patterns are not well characterized. For example, the molecule encoded by B*5306 exhibits both B5 and B53 reactivity [7]. Table 8 provides a description of some of these variant reaction patterns for HLA class I alleleic products. A number of officially named HLA specificities cannot readily be identified by the majority of typing laboratories. Both NMDP and the UCLA cell exchange data demonstrate that relatively new specificities such as A203, A2403, B5102, and B703, are rarely used. They are usually designated by their associated or broad specificities (A2, A24, B51, and B7, respectively). Even specificities such as A66, B64, and B65 are poorly identified. Therefore, the percentage recognition in the included tables can be helpful in selecting alternative antigens as search determinants.

HLA Alleles and Serologic Types

841

TABLE 5 HLA–DRB1 alleles and their serologic designations International cell exchange UCLA HLA allele

WHO assigned

Cells tested

DRB1*0101 DRB1*0102 DRB1*0103 DRB1*0104 DRB1*0105 DRB1*0106 DRB1*0107 DRB1*0301 DRB1*0302 DRB1*0303 DRB1*0304 DRB1*0305 DRB1*0306 DRB1*0307 DRB1*0308 DRB1*0309 DRB1*0310 DRB1*0311 DRB1*0312 DRB1*0313 DRB1*0314 DRB1*0315 DRB1*0316 DRB1*0317 DRB1*0318 DRB1*0401 DRB1*0402 DRB1*0403 DRB1*0404 DRB1*0405 DRB1*0406 DRB1*0407 DRB1*0408 DRB1*0409 DRB1*0410 DRB1*0411 DRB1*0412 DRB1*0413 DRB1*0414 DRB1*0415 DRB1*0416 DRB1*0417 DRB1*0418 DRB1*0419 DRB1*0420 DRB1*0421 DRB1*0422 DRB1*0423 DRB1*0424 DRB1*0425 DRB1*0426 DRB1*0427 DRB1*0428 DRB1*0429 DRB1*0430 DRB1*0431 DRB1*0432 DRB1*0433 DRB1*0434 DRB1*0435

DR1 DR1 DR103 DR1 – – – DR17(3) DR18(3) DR18(3) DR17(3) DR17(3) DR3 DR3 – – DR17(3) DR17(3) DR3 – – – – – – DR4 DR4 DR4 DR4 DR4 DR4 DR4 DR4 DR4 DR4 DR4 – DR4 DR4 DR4 DR4 DR4 – DR4 DR4 DR4 DR4 DR4 DR4 DR4 DR4 – DR4 DR4 – DR4 DR4 – – –

12 2 1 – – – – 22 10 – 1 – – – – – – – – – – – – – – 5 4 7 12 9 2 2 – – – 1 – – – 2 1 – – – – – – – – – 2 – – – – – 1 – – –

Assigned type (%) DR1 [88–100] DR1 [86–95] DR103 [70] DR1 [17] – – – – DR17 [59–69] DR3 [28–41] DR18 [62–67] DR3 [27–35] – DR3 [51] DR17 [47] – – – – – – – – – – – – – – DR4 [100] DR4 [84–100] DR4 [97–100] DR4 [95–100] DR4 [94–100] DR4 [89–96] DR4 [97–98] – – – DR4 [98] – – – DR4 [84–92] DR11 [24–49] DR4 [100] – – – – – – – – – DR4 [100] – – – – – DR4 [100] – – –

NMDP Cells tested

Assigned type (%)

9538 2459 1666 1 1 – – 12867 370 7 6 – 2 – – – 2 2 – – – – – – – 10271 1178 960 3003 591 84 1154 418 16 28 74 1 6 2 4 1 1 – – – 1 – 1 – – – – – – – – – – – –

DR1 [97] DR1 [95] DR1 [42] DR103 [33] blank [15] DR1 [100] @ DR1 [100] @ – – DR17 [61] DR3 [36] DR18 [63] DR3 [28] DR17 [2] DR18 [45] DR17 [29] DR3 [14] DR3 [67] DR17 [17] – DR3 [50] blank [50] @ – – – DR3 [50] DR17 [50] @ DR3 [50] DR17 [50] @ – – – – – – – DR4 [97] DR4 [91] DR4 [94] DR4 [92] DR4 [94] DR4 [91] DR4 [90] DR4 [89] DR4 [94] DR4 [86] DR4 [90] DR4 [100] @ DR4 [69] DR4 [100] @ DR4 [75]@ DR4 [100] @ DR4 [100] @ – – – DR4 [100] @ – § – – – – – – – – – – – –

Comments

DR3†

[18]

DR4⫻11

DR4⫻3 DR4short

[18]

842

G. M. Th. Schreuder et al.

TABLE 5 (Continued) International cell exchange UCLA HLA allele

WHO assigned

Cells tested

DRB1*0436 DRB1*0437 DRB1*0438 DRB1*0701 DRB1*0703 DRB1*0704 DRB1*0801 DRB1*0802 DRB1*0803 DRB1*0804 DRB1*0805 DRB1*0806 DRB1*0807 DRB1*0808 DRB1*0809 DRB1*0810 DRB1*0811 DRB1*0812 DRB1*0813 DRB1*0814 DRB1*0815 DRB1*0816 DRB1*0817 DRB1*0818 DRB1*0819 DRB1*0820 DRB1*0821 DRB1*0822 DRB1*0823 DRB1*0901 DRB1*1001 DRB1*1101 DRB1*1102 DRB1*1103 DRB1*1104 DRB1*1105 DRB1*1106 DRB1*1107 DRB1*1108 DRB1*1109 DRB1*1110 DRB1*1111

– – – DR7 DR7 DR7 DR8 DR8 DR8 DR8 DR8 DR8 DR8 – DR8 DR8 DR8 DR8 – DR8 – DR8 DR8 – – – – – – DR9 DR10 DR11(5) DR11(5) DR11(5) DR11(5) DR11(5) DR11(5) DR11(5) DR11(5) DR11(5) DR11(5) DR11(5)

– – – 15 – – 4 1 11 7 – 1 – – – – – – – – – – – – – – – – – 16 11 8 5 2 11 – – – – – – 2

DRB1*1112 DRB1*1113 DRB1*1114 DRB1*1115 DRB1*1116

– DR11(5) DR11(5) – –

– – – – –

– – – DR7 [97–100] – – DR8 [95–100] DR8 [99] DR8 [97–100] DR8 [88–99] – DR8 [100] – – – – – – – – – – – – – – – – – DR9 [97–100] DR10 [94–95] DR11 [95–100] DR11 [89–100] DR11 [84–100] DR11 [94–100] – – – – – – DR11 [87–90] DR13 [7–8] DR5 [4–5] – – – – –

DRB1*1117 DRB1*1118 DRB1*1119 DRB1*1120 DRB1*1121 DRB1*1122 DRB1*1123 DRB1*1124 DRB1*1125 DRB1*1126 DRB1*1127

– – DR11(5) DR11(5) DR11(5) – DR11(5) – DR11(5) DR11(5) DR11(5)

– – 1 – 1 – – – – – –

– – DR11 [100] – DR11 [97] – – – – – –

Assigned type (%)

NMDP Cells tested – – – 13382 – – 1975 314 285 241 11 21 4 – 1 2 9 – – – – – – – – – – – – 1869 1490 4934 527 553 2622 5 16 3 – 5 2 7

Assigned type (%) – – – DR7 [99] – – DR8 [96] DR8 [97] DR8 [90] DR12 DR8 [95] DR8 [100] DR8 [86] DR8 [100] @ – DR8 [100]@ DR8 [100]@ DR8 [89] – – – – – – – – – – – – DR9 [97] DR10 [93] DR1 DR11 [93] DR5 DR11 [89] DR5 DR11 [91] DR5 DR11 [89] DR5 DR11 [81] DR5 [69] DR11 DR11 [100] @ – DR11 [80] DR5 DR11 [52]@ DR11 [78]

Comments

[19] [3] blank [4]

DR8var

[2] blank [2] [4] [3] DR6 [3] [2] DR13 [2] [4] [25] DR11⫻3 [20]@

– 5 3 1 –

– DR11 [80] DR14 [20]@ DR11 [100]@ DR11 [100]@ –

1 1 2 – 1 – – 2 – – –

DR14 § DR11 – DR11 – – DR11 – – –

DR11⫻13 DR11⫻14 DR11 [70] DR13 [30]$ DR11⫻13

[100]@ [100]@ DR11⫻13 [100]@ [50] blank [50]@

HLA Alleles and Serologic Types

843

TABLE 5 (Continued) International cell exchange UCLA HLA allele

WHO assigned

Cells tested

DRB1*1128 DRB1*1129 DRB1*1130 DRB1*1131 DRB1*1132 DRB1*1133 DRB1*1134 DRB1*1135 DRB1*1136 DRB1*1137 DRB1*1138 DRB1*1139 DRB1*1140 DRB1*1141 DRB1*1201 DRB1*1202 DRB1*1203 DRB1*1204 DRB1*1205 DRB1*1206 DRB1*1207 DRB1*1301 DRB1*1302 DRB1*1303

– DR11(5) – – – – – – – – – – – – DR12(5) DR12(5) DR12(5) DR5 DR12(5) DR12(5) – DR13(6) DR13(6) DR13(6)

– – – – – – – – – – – – – – 8 15 – – – – – 8 15 10

DRB1*1304

DR13(6)

–

DRB1*1305

DR13(6)

3

DRB1*1306 DRB1*1307 DRB1*1308 DRB1*1309 DRB1*1310 DRB1*1311 DRB1*1312

DR13(6) DR13(6) DR13(6) – DR13(6) DR13(6) DR6

– – – – 2 – –

DR13 [48–62] DR11 [11–34] DR6 [5–20] DR14 [3–8] – – – – DR13 [84–85] DR6 [7–15] – –

DRB1*1313 DRB1*1314 DRB1*1315 DRB1*1316 DRB1*1317 DRB1*1318 DRB1*1319 DRB1*1320 DRB1*1321 DRB1*1322 DRB1*1323 DRB1*1324 DRB1*1325 DRB1*1326 DRB1*1327 DRB1*1328 DRB1*1329 DRB1*1330 DRB1*1331 DRB1*1332 DRB1*1333 DRB1*1334

DR13(6) DR13(6) – DR13(6) DR13(6) DR13(6) DR13(6) DR13(6) – DR13(6) – – – – DR13(6) – DR6 – – – – –

1 – – – – – – – – – – – – – – – – – – – – –

DR13 [83] DR6 [13] – – – – – – – – – – – – – – – – – – – – –

Assigned type (%) – – – – – – – – – – – – – – DR12 [85–96] DR13 [5–12] DR12 [97–100] – – – – – DR13 [88–90] DR6 [3–8] DR13 [85–97] DR6 [7–9] DR13 [50–76] DR6 [10–21] DR‘1303’ [0–11] DR14 [0–17] DR3 [0–8] –

NMDP Cells tested 1 – – – – – 1 – – – – – – – 1399 303 4 – – 2 – 5867 4646 1394 139 363 8 2 11 1 9 – 8 – 1 1 – – – – 1 1 3 – – – – – – – – – – – –

Assigned type (%) DR11 [100]@ – – – – – DR11 [100]@ – – – – – – – DR12 [88] DR5 [3] DR11 [2] DR12 [95] DR12 [100]@ – – DR12 [100]@ – DR13 [84] DR6 [13] DR13 [78] DR6 [13] DR14 [1] DR13 [75] DR6 [8] DR5 [2] DR11 [2] DR14 [2] DR14 [2] blank [4] DR13 [86] DR5 [4] DR6 DR6 [2] blank [4] DR13 [61] DR6 [11] DR11 [7] DR12 [5] DR5 [3] DR14 [1] DR13 [72] DR6 [25] DR13 [100]@ DR13 [47] DR6 [53] § DR13 [100] – DR13 [37] DR11 [25] DR12 [13] DR6 [13] – DR11 [100]@ § – – – – DR13 [100]@ DR13 [100]@ DR13 [100]@ – – – – – – – – – – – –

Comments

DR12⫻11

DR13⫻12 DR52–neg (18)

DR13⫻8

DR13 [100]$

DRblank–DR52pos

844

G. M. Th. Schreuder et al.

TABLE 5 (Continued) International cell exchange UCLA HLA allele

WHO assigned

Cells tested

DRB1*1335 DRB1*1336 DRB1*1337 DRB1*1338 DRB1*1339 DRB1*1340 DRB1*1341 DRB1*1342 DRB1*1343 DRB1*1344 DRB1*1345 DRB1*1346 DRB1*1347 DRB1*1401 DRB1*1402

– DR13(6) – – – – – DR13(6) – – – – – DR14(6) DR14(6)

– – – – – – – – – – – – – 15 6

DRB1*1403 DRB1*1404

DR1403 DR1404

– 8

DRB1*1405

DR14(6)

2

DRB1*1406

DR14(6)

1

DRB1*1407 DRB1*1408 DRB1*1409 DRB1*1410 DRB1*1411

DR14(6) DR14(6) – DR14(6) DR14(6)

– 1 – 1 1

DRB1*1412 DRB1*1413 DRB1*1414 DRB1*1415 DRB1*1416 DRB1*1417 DRB1*1418 DRB1*1419 DRB1*1420 DRB1*1421

DR14(6) DR14(6) DR14(6) DR8 DR6 DR6 DR6 DR14(6) DR14(6) DR6

– 1 1 1 – – – – – 1

DRB1*1422 DRB1*1423 DRB1*1424 DRB1*1425 DRB1*1426 DRB1*1427 DRB1*1428 DRB1*1429 DRB1*1430 DRB1*1431 DRB1*1432 DRB1*1433 DRB1*1434 DRB1*1435 DRB1*1436 DRB1*1437 DRB1*1438 DRB1*1439 DRB1*1440 DRB1*1501

DR14(6) – – – DR14(6) DR14(6) – DR14(6) – – – – – – – – – – – DR15(2)

– – – – – – – – – – – – – – – – – – – 19

Assigned type (%) – – – – – – – – – – – – – DR14 [90–92] DR6 [5–7] DR14 [46–75] DR6 [12–23] DR17 [8–15] DR13 [0–8] – DR14 [84–89] DR1404 [5–8] DR6 [3–7] DR8 [0–10] DR14 [74–85] DR6 [11–15] DR3 [0–13] DR13 [28] DR14 [26] DR6 [19] DR3 [11] – DR14 [94] – DR14 [76] DR6 [10] DR14 [59] DR6 [12] DR11 [12] DR‘14⫻11’ [9] – DR14 [59] DR6 [17] DR13 [10] DR14 [89] DR13 [5] DR8 [50] DR14 [26] DR13 [8] – – – – – DR14 [49] DR13 [23] DR6 [19] DR17 [6] – – – – – – – – – – – – – – – – – – – DR15 [78–91] DR2 [7–22]

NMDP Cells tested – – – – – – – – – – – – – 2654 234 46 116 63 130 15 1 – 5 2 – – – – 4 1 – – 1 1 – – – – – – – – – – – – – – – – – – – 9701

Assigned type (%)

Comments

– – – – – – – – – – – – – DR14 [78] DR6 [13] DR13 [2] DR14 [65] DR6 [11] DR13 [9] DR5 [6] blank [6] DR14 [66] DR6 [12] DR13 [7] DR14 [73] DR6 [9] DR1404 [7] DR12 [2] DR8 [2] DR14 [84] DR6 [11] DR13 [3] DR14 [46] DR6 [15] DR13 [13] DR5 [11] blank [8] DR14 [93] DR14 [100]@ – DR14 [100]@ DR14 [100]@ – – – – DR14 [75] DR13 [25]@ DR14 [100]@ – – § DR6 [100]@ – – – – – – – – – – – – – – – – – – – DR15 [76] DR2 [22]

DR14† DR14⫻11

DR8⫹DR52 DR14⫻13 DR14⫻13 DR14–DQ7† DR13†

HLA Alleles and Serologic Types

845

TABLE 5 (Continued) International cell exchange UCLA HLA allele

WHO assigned

Cells tested

DRB1*1502 DRB1*1503 DRB1*1504 DRB1*1505 DRB1*1506 DRB1*1507 DRB1*1508 DRB1*1509 DRB1*1510 DRB1*1511 DRB1*1601 DRB1*1602

DR15(2) DR15(2) DR15(2) DR15(2) DR15(2) DR15(2) DR2 – – – DR16(2) DR16(2)

11 6 – – – – – – – – 1 6

DRB1*1603 DRB1*1604 DRB1*1605 DRB1*1607 DRB1*1608

DR2 DR16(2) DR2 – –

– – – – –

Assigned type (%) DR15 [87–90] DR2 [7–10] DR15 [86–94] DR2 [3–10] – – – – – – – – DR16 [31] DR15 [38] DR2 [31] DR16 [63–85] DR2 [11–17] DR15 [7–8] – – – – –

NMDP Cells tested 816 352 1 2 1 – – – – – 1150 290 – 2 1 – –

Assigned type (%) DR15 DR15 DR15 DR15 DR15 – – – – – DR16 DR16

Comments

[75] DR2 [18] DR16 [1] [84] DR2 [11] [100]@ [50]@ [100]@ (19)

[51] DR2 [27] DR15 [13] [54] DR2 [26] DR15 [15]

– DR16 [100]@ DR16 [100]@ – –

DR16 †

@ ⫽ allele has been reported ⬍6 times and/or serologically identified in ⬍4 individuals. § ⫽ allele identified but serology not informative. $ ⫽ HLA-Club Cell Exchange: one sample typed by 17 laboratories and allele identified by at least three laboratories. † ⫽ locally identified in Leiden.

TABLE 6 HLA-DRB3/4/5 alleles and their serologic designations

HLA alleles DRB3*0101 DRB3*0102 DRB3*0103 DRB3*0104 DRB3*0105 DRB3*0106 DRB3*0107 DRB3*0201 DRB3*0202 DRB3*0203 DRB3*0204 DRB3*0205 DRB3*0206 DRB3*0207 DRB3*0208 DRB3*0209 DRB3*0210 DRB3*0211 DRB3*0212 DRB3*0213 DRB3*0301 DRB3*0302

International cell exchange WHO Cells UCLA assigned assigned tested type (%) Comments DR52 DR52 – – – DR52 DR52 DR52 DR52 DR52 – – – DR52 DR52 DR52 DR52 DR52 – – DR52 DR52

53 1 – – – – – 2 72 1 – – – – – – – – – – 28 –

DR52 DR52 – – – – – DR52 DR52 DR52 – – – – – – – – – – DR52 –

[91–100] [99]

[97–99] [89–100] [100]

[85–100]

TABLE 6 (Continued)

HLA alleles DRB3*0303 DRB4*0101 DRB4*0102 DRB4*0103 DRB4*0103102N DRB4*0104 DRB4*0105 DRB4*0201N DRB4*0301N DRB5*0101 DRB5*0102 DRB5*0103 DRB5*0104 DRB5*0105 DRB5*0106 DRB5*0107 DRB5*0108N DRB5*0109 DRB5*0110N DRB5*0202 DRB5*0203 DRB5*0204 DRB5*0205

International cell exchange WHO Cells UCLA assigned assigned tested type (%) Comments – DR53 DR53 DR53 Null – DR53 Null Null DR51 DR51 – – – – DR51 Null – Null DR51 – – –

– 33 – 28 7 – – – – 22 6 – – – – – – – – 6 – – –

– DR53 – DR53 DR53 – – – – DR51 DR51 – – – – – – – – DR51 – – –

[100] [90–100] [42–100]

[82–87] [86–89]

[86–92]

846

G. M. Th. Schreuder et al.

TABLE 7 HLA-DQB1 alleles and their serologic designations International cell exchange UCLA HLA allele

WHO Cells assigned tested

DQB1*0201 DQB1*0202 DQB1*0203 DQB1*0301

DQ2 DQ2 DQ2 DQ7(3)

7 5 – 21

DQB1*0302

DQ8(3)

13

DQB1*0303

DQ9(3)

4

DQB1*0304 DQB1*0305 DQB1*0306 DQB1*0307 DQB1*0308 DQB1*0309 DQB1*0310 DQB1*0401 DQB1*0402 DQB1*0501 DQB1*0502

DQ7(3) DQ8(3) DQ3 – – – DQ8(3) DQ4 DQ4 DQ5(1) DQ5(1)

1 – – – – – – 1 7 10 7

DQB1*0503 DQB1*0504 DQB1*0601 DQB1*0602 DQB1*0603 DQB1*0604 DQB1*0605 DQB1*0606 DQB1*0607 DQB1*0608 DQB1*0609 DQB1*0610 DQB1*0611 DQB1*0612 DQB1*0613 DQB1*0614 DQB1*0615 DQB1*0616 DQB1*0617

DQ5(1) DQ5(1) DQ6(1) DQ6(1) DQ6(1) DQ6(1) DQ6(1) – – – DQ6(1) – DQ1 DQ1 – DQ6(1) – – –

3 1 2 7 3 1 – – – – 1 – – – – – – – –

Assigned type (%) DQ2 [93–100] DQ2 [90–94] – DQ7 [70–80] DQ3 [16–27] DQ8 [0–8] DQ3 [47–74] DQ8 [22–56] DQ7 [3–7] DQ9 [0–6] DQ3 [45–60] DQ9 [35–42] DQ7 [60] DQ3 [34] – – – – – – DQ4 [88] DQ4 [81–91] DQ1 [51–71] DQ5 [29–41] DQ6 DQ1 [54–64] DQ6 [13–27] DQ5 [0–24] DQ1 [59] DQ6 [24] DQ5 [14] DQ1 [66] DQ5 [23] DQ6 [8] DQ1 [49–54] DQ6 [40] DQ1 [48–57] DQ6 [38–48] DQ1 [42–58] DQ6 [34–56] DQ1 [50] DQ6 [44] – – – – DQ6 [54] DQ1 [43] – – – – – – – –

NMDP Cells tested 10258 882 2 14154

Assigned type (%) DQ2 DQ2 DQ2 DQ7

Comments

[97] [81] DQ1 [6] [50]@ [81] DQ3 [12] DQ8 [1]

7814 DQ8 [62] DQ3 [28] DQ7 [3] 3110 DQ9 [64] DQ3 [16] DQ2 [10] DQ8 [1] DQ7 [1] 120 DQ3 [44] DQ7 [37] DQ8 [8] 60 DQ8 [36] DQ3 [34] 1 DQ8 [100]@ – – – – – – – – 124 DQ4 [77] DQ3 [7] DQ8 [4] DQ7 [3] 1197 DQ4 [87] DQ3 [3] blank [6] 6537 DQ5 [69] DQ1 [20] DQ6 [1] 997 DQ5 [50] DQ1 [24] DQ6 [14] 1540 63 674 6390 3587 2065 379 1 4 8 150 – – – 1 – – – –

DQ5 DQ5 DQ6 DQ6 DQ6 DQ6 DQ6 DQ5 DQ1 DQ6 DQ6 – – – § – – – –

[60] DQ1 [55] DQ1 [66] DQ1 [67] DQ1 [62] DQ1 [60] DQ1 [76] DQ1 [100]@ [100]@ [39] DQ1 [47] DQ1

[22] [23] [23] [26] [23] [25] [15]

DQ7negDQ8neg#

DQ6 [4] DQ6 [2] DQ5 [1] DQ5 [2] DQ5 [2] DQ5 [1]

[28] [36] DQ5 [2]

DQ6 †

@ ⫽ allele has been reported ⬍6 times and/or serologically identified in ⬍4 individuals. § ⫽ allele identified but serology not informative. † ⫽ locally identified in Leiden.

FUTURE PLANS For many alleles, serologic equivalents have now been identified and represent between 64 and 79% of the WHO designated alleles (Table 1). It will be important to include samples with poorly defined serologic types into serologic testing programs such as the International Cell Exchange (UCLA) or other locally organized cell exchange programs. Laboratories are also encouraged to participate in the 13th International Histocompatibility Workshop serology component, which will study rare and newly identified HLA-A and -B alleles

by serological methods [8]. Therefore, laboratories that encounter rare or poorly identified alleles during typing procedures, should store typing material, preferably viable lymphocytes or B cell lines for further testing. CONCLUSION This report presents a database listing WHO HLA antigen nomenclature and serologic equivalents of 123 HLA-A, 272 HLA-B, and 155 HLA-DRB1 alleles. The

HLA Alleles and Serologic Types

847

TABLE 8 HLA class I alleles expressed as HLA-A and -B variants for which no official nomenclature is available and/or with known variation in serologic reaction pattern. Allele

Serotype

A*1104 A*1105 A*2408 A*2416 A*2418 A*2419 A*2421 A*2608 A*2610 A*3204 A*3404 A*6603 A*6812 B*0703 B*0708 B*0715 B*0716 B*0720 B*0724 B*0802 B*0803 B*0804 B*0810 B*1303 B*1304

B*2715 B*3515 B*3531 B*3702 B*3803 B*3905 B*4008 B*4010 B*4012 B*4019 B*4021

A11short A11var A24var A31var A24⫻3 A9short A9var A10short A10var A3 A34x31 A10short A28short B703, BPOT B7short B7(Bw6var) B7short B7short B7weak B8var(Bw4⫹) B8var(Bw4⫹) B8neg(Bw6⫹) B8var(Bw6⫹) B21var B15x21 (Bw4⫹) ‘NM5’ B62(Bw4⫹) B15short B62var B18(Bw6⫺) B18(Bw4⫹) B2708, B7Qui B27var B’X’(Bw6⫹), B40var B’X’(Bw6⫹) B35var Bfu-var B‘37x27’ B16(Bw4⫹) B39var B40x48var B60var B48x70 B40(Bw4⫹) B40x15

B*4025 B*4406 B*4408

B40short B12var/’blank’ B44short

B*4409 B*4415 B*4416 B*4602 B*4702

B12(Bw6⫹) B44var B47/44 B46var B47(Bw6⫹)

B*1523 B*1524 B*1537 B*1538 B*1806 B*1809 B*2708 B*2711 B*2712

Description of serological pattern Reactive with A11, A11⫹3(⫹1) sera; negative with A11⫹6601 sera Only 3 out of 31 A11 antisera reactive Similar to A2403; A24 sera positive, A9 sera negative A31-like. A*3101 with 3⬘ ex 2 partly homologous to A*24 alleles Reactive with most A3, A9 and A24 sera Very short A9, shorter than A2403 A9 sera positive, A24 and A23 sera negative A26 and A66 specific sera negative A26 short A19 and A32 sera neg. All A3 sera positive. One A32 mAb pos Reactive with A34 and some of the A31 sera A26 and A66 specific sera negative Most A28⫹2 sera negative, A28⫹33⫹34⫹sera positive Short B7 with B7⫹42 sera positive, but B7⫹22 sera negative Very few B7 sera reactive; similar to B*0703 B7 with aberrant Bw6 epitope. Some Bw6 sera non-reactive Very few B7 sera reactive; pattern similar to B703 B7⫹42 sera reactive; B7 specific sera weak/negative B7 sera weak, B7⫹27(⫹40) sera positive Reactive with Bw4, B8 and B8⫹59 sera; negative with Bw6 sera

Reference

[20]

[20] [11] 2 3

[11] [21] 4

[22] [23] [30] [24] [5]

Some B8⫹59 and B49 sera positive; most B8 sera negative Short B8; part of B8 specific sera non-reactive B21 like specificity; also some B15 sera reactive; Bw4 positive Similar to B*1303; Some B15 and B21 sera reactive; UCLA Int. cell exchange TER 847; HLA-Club cell exchange B5/B53/B77; B5 CREG sera positive; B70-Bw4 Regular B15 and B62 sera positive; Bw6 negative, Bw4 positive Very few B15 sera reactive, also typed as B70; Bw6 associated B62⫹B52, B62, B15 and B52 sera positive B18 which is non-reactive with Bw4 and Bw6 sera B18 with Bw4 epitope Most B7 sera positive, but B27 sera mostly negative; Bw6 associated B27 with some B40 sera positive Reactive with few B40 and B27⫹7, B7CREG sera; B27 and B7 specific sera negative B27 sera negative, reactive with some Bw4 sera Only part of B35 sera reactive. B75 CREG sera reactive Bfu-B40 like pattern Some B37 and some B27 sera reactive; Bw4 associated Some B16 sera reactive; B38 and B39 sera negative Previously called ST-16; Some B38 reactivity and some B16/B39 sera non-reactive Only some B40⫹48(⫹7) CREG sera reactive; tested on TER-969 Some B60 sera reactive; B48-like Additional to B40⫹48 also some B15 ⫹B70 CREG sera positive B*4002 sequence with Bw4 epitope Reactive with B40⫹48, B15⫹57 and some other B15 sera; neg with B40⫹13⫹47 sera Only 2 out of 15 B40 sera reactive Few B12 sera reactive; can easily be missed Reactive with all B44, most B12 and most B62⫹63⫹57 sera; Detected in Welsh population but extremely rare B*4402 allele with Bw6 sequence; will be typed as B45 B*4501 allele with Bw4 sequence: will be typed as B44 B47 sera positive, few B12 and B44 sera positive; tested on TER-1017 B46⫹75 sera negative, B46⫹Cw3 sera reactive Reactive with B40⫹47⫹13 sera; B27⫹47 sera negative

[25] [11] [11]

[22]

4

[31]1 [9]

[26] [16] [27] [10]

[15] [28] 4

848

G. M. Th. Schreuder et al.

TABLE 8 (Continued) Allele

Serotype

B*4703

B47var

B*4802 B*5106 B*5112 B*5304 B*5306 B*5307

B48? B5 B‘blank’ B52⫻53 B51⫻53 B53⫻37

B*5503 B*5509 B*5603

B55⫻B67 var B22short B22var

B*5604 B*5606 B*5607 B*8201

B56var B78var B56(Bw4⫹) B45x56 var

1 2 3 4

⫽ ⫽ ⫽ ⫽

Description of serological pattern

Reference

Strongly reactive with B40⫹47⫹13, but not with B27⫹47 sera; aberrant Bw4 epitope, reactive with both Bw4 and Bw6 antisera; tested on TER977 B70⫺B71-like B51 specific sera negative Possibly not expressed B5⫹53 sera reactive B51 and B53 specific sera positive, some B5 sera negative B53 with mAb 12W069 (anti B37⫹52⫹18⫹39) positive, but most allo B37 sera negative Maybe less reactive with Bw6 sera; Cw1 associated B55 and B56 specific sera negative; only few broad B22 sera reactive B22 with some B15 features; reactive with B22 sera and with Bw6; detected in Orientals Reactive with part of B56 and B22 specific sera; infrequent in Korean population Some B78 sera positive, all B22 sera negative Bw56 with Bw4 epitope Some B22 and B45 specific sera reactive; B12 sera negative

[29] [11] [10] [7] 4

1

[7] [22] [11]

E. M. van den Berg-Loonen, personal communication. A. de Smet, Antwerp, Belgium, personal communication. HLA-Club Cell Exchange: one sample typed by serology in 17 laboratories. local observation in Leiden.

dictionary is an update of those published in 1997 [1] and 1999 [2]. The data summarize information obtained through the International Cell Exchange (UCLA), the National Marrow Donor Program (NMDP) and by individual laboratories. In addition, a listing is provided of alleles which are expressed as antigens with serologic reaction patterns that differ from the well established HLA specificities and that often lack official WHO nomenclature designations. The provided equivalents will be useful in guiding searches for unrelated tissue donors in which patient and/or potential donors are typed by either serology or by DNA-based methods. ACKNOWLEDGMENTS

We thank Professor Dr. E. M. van den Berg-Loonen (Maastricht, The Netherlands) and Dr. C. Darke (Cardiff, UK) for their personal contributions. The efforts from the participants of the UCLA International Cell Exchange are gratefully acknowledged. We specifically thank the typing laboratories in the Netherlands, Belgium, and Luxembourg which participate in the HLA-Club Cell Exchange, for submitting, typing, and commenting on the samples with new and rare HLA alleles. This work was supported in part by the J.A. Cohen Institute for Radiopathology and Radioprotection (IRS), by the Dutch National Reference Center for Histocompatibility, and by the U.S. Office of Naval Research N00014-96-2-0016, N0001499-2-0006, to the National Marrow Donor Program and N00014-99-1-0551 to the C.W. Bill Young Marrow Donor Recruitment and Research Program.

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