Nomenclature for Factors of the HLA System
July 2013
Compiled by Steven G. E. Marsh for the WHO Nomenclature Committee for Factors of the HLA System.
Published In:
- Tissue Antigens. 2013 82:447-51
- Human Immunology. 2013 74:1709-12
- International Journal of Immunogenetics. 2013 40:528-32
The following sequences have been submitted to the Nomenclature Committee since the June 2013 nomenclature update and, following agreed policy, have been assigned official allele designations. Full details of all sequences will be published in a forthcoming report.
Below are listed the newly assigned sequences (Table 1) and confirmations of previously reported sequences (Table 2). The accession number of each sequence is given and these can be used to retrieve the sequence files from the EMBL, GenBank or DDBJ data libraries. Although accession numbers have been assigned by the data-libraries and most sequences are already available, there is still the possibility that an author may not yet have allowed the sequence to be released; in such a case you will have to contact the submitting author directly. Additional information pertaining to new sequences is often included in the publications describing these alleles; a listing of recent publications that describe new HLA sequences is given in Table 3.
All new and confirmatory sequences should now be submitted directly to the WHO Nomenclature Committee for Factors of the HLA System via the IMGT/HLA Database using the sequence submission tool provided (2). The IMGT/HLA Database may be accessed via the world wide web at : http://ww.ebi.ac.uk/imgt/hla/.
Table 1: New Sequences
| Sequence | Cell identification | Accession number | Submitting author |
|---|---|---|---|
| A*02:01:96 | 90185817 | HG324057 | Christian Seidl, Frankfurt am Main, Germany |
| A*02:11:05 | HG00005973 | KC875889 | Histogenetics, Ossining, USA |
| A*24:02:71 | CALE747852 | HG007959 | Steven GE Marsh, London, United Kingdom |
| A*24:247 | SZGSQ 21 | KF304799 | Su-Qing Gao, Shenzhen, China |
| A*26:02:02 | JMDP36K0199 | AB829894 | Kenji Tadokoro, Tokyo, Japan |
| A*33:72 | BY01088 | KF278711 | Carolyn K Hurley, Washington DC, USA |
| A*68:103 | BY01084 | KF278715 | Carolyn K Hurley, Washington DC, USA |
| A*68:104 | BY01086 | KF278713 | Carolyn K Hurley, Washington DC, USA |
| B*07:199 | BY01056 | KF266789 | Carolyn K Hurley, Washington DC, USA |
| B*08:01:22 | HG00005989 | KC991056 | Histogenetics, Ossining, USA |
| B*08:112 | HG00005988 | KC748294 | Histogenetics, Ossining, USA |
| B*13:71 | HG00005990 | KC875907 | Histogenetics, Ossining, USA |
| B*15:283 | BY01082 | KF266791 | Carolyn K Hurley, Washington DC, USA |
| B*18:92 | HG00005980 | KF015527 | Histogenetics, Ossining, USA |
| B*18:93 | HG00005984 | KC959276 | Histogenetics, Ossining, USA |
| B*18:94N | HG00005985 | KC603580 | Histogenetics, Ossining, USA |
| B*27:05:23 | HG00005977 | KC959277 | Histogenetics, Ossining, USA |
| B*27:05:24 | HG00005979 | KC991055 | Histogenetics, Ossining, USA |
| B*27:07:04 | HG00005978 | KC959291 | Histogenetics, Ossining, USA |
| B*35:234 | HG00005993 | KC904411 | Histogenetics, Ossining, USA |
| B*35:235 | HG00005994 | KC904405 | Histogenetics, Ossining, USA |
| B*35:236 | HG00005996 | KC904403 | Histogenetics, Ossining, USA |
| B*35:237 | HG00005997 | KC904409 | Histogenetics, Ossining, USA |
| B*35:238 | HG00005998 | KC959289 | Histogenetics, Ossining, USA |
| B*35:239 | BY01076 | KF266797 | Carolyn K Hurley, Washington DC, USA |
| B*35:240 | IME956-10T | KF267447 | Manuela Testi, Rome, Italy |
| B*35:241 | 35904 | HF970578 | Marcel GJ Tilanus, Maastricht, The Netherlands |
| B*37:38 | HG00005981 | KC959281 | Histogenetics, Ossining, USA |
| B*37:39 | HG00005982 | KC875900 | Histogenetics, Ossining, USA |
| B*37:40 | HG00005983 | KC875910 | Histogenetics, Ossining, USA |
| B*37:41 | HG00005987 | KC904401 | Histogenetics, Ossining, USA |
| B*40:247 | 101762 | KF208638 | Chun-Ying Lin, Taipei, Taiwan |
| B*40:248 | BY01075 | KF266798 | Carolyn K Hurley, Washington DC, USA |
| B*41:29 | HG00005976 | KC959288 | Histogenetics, Ossining, USA |
| B*48:32 | BY01071 | KF266802 | Carolyn K Hurley, Washington DC, USA |
| B*50:34 | BY01074 | KF266799 | Carolyn K Hurley, Washington DC, USA |
| B*52:01:18 | HG00005999 | KC904419 | Histogenetics, Ossining, USA |
| B*52:32 | BY01081 | KF266792 | Carolyn K Hurley, Washington DC, USA |
| B*57:66 | BY01072 | KF266801 | Carolyn K Hurley, Washington DC, USA |
| B*58:01:14 | HG00005992 | KC959279 | Histogenetics, Ossining, USA |
| B*58:47 | HG00005991 | KC875906 | Histogenetics, Ossining, USA |
| C*01:02:20 | D10047948 | KF175394 | Jinghui Qu, Shenzhen, China |
| C*01:02:21 | D10057079 | KF175397 | Jinghui Qu, Shenzhen, China |
| C*01:02:22 | D10097769 | KF175399 | Jinghui Qu, Shenzhen, China |
| C*01:02:23 | D10092834 | KF175403 | Jinghui Qu, Shenzhen, China |
| C*01:79:02 | D10098667 | KF175396 | Jinghui Qu, Shenzhen, China |
| C*01:80 | D10066432 | KF175395 | Jinghui Qu, Shenzhen, China |
| C*01:81 | D10096758 | KF175398 | Jinghui Qu, Shenzhen, China |
| C*01:82 | D10111537 | KF175400 | Jinghui Qu, Shenzhen, China |
| C*01:83 | D10128630 | KF175401 | Jinghui Qu, Shenzhen, China |
| C*01:84 | D10075614 | KF175402 | Jinghui Qu, Shenzhen, China |
| C*02:69 | 35767 | HF674389 | Marcel GJ Tilanus, Maastricht, The Netherlands |
| C*02:70 | D10050477 | KF175406 | Jinghui Qu, Shenzhen, China |
| C*02:71 | D10028882 | KF175404 | Jinghui Qu, Shenzhen, China |
| C*02:72 | D10025544 | KF175405 | Jinghui Qu, Shenzhen, China |
| C*03:03:22 | D10070263 | KF175415 | Jinghui Qu, Shenzhen, China |
| C*03:04:37 | D10064091 | KF175421 | Jinghui Qu, Shenzhen, China |
| C*03:04:38 | D10116860 | KF175422 | Jinghui Qu, Shenzhen, China |
| C*03:04:39 | D10074696 | KF175424 | Jinghui Qu, Shenzhen, China |
| C*03:04:40 | D10085847 | KF175426 | Jinghui Qu, Shenzhen, China |
| C*03:04:41 | D10047135 | KF175427 | Jinghui Qu, Shenzhen, China |
| C*03:13:02 | D10021653 | KF175431 | Jinghui Qu, Shenzhen, China |
| C*03:196 | JMDP36K0161 | AB829605 | Kenji Tadokoro, Tokyo, Japan |
| C*03:197 | D10098824 | KF175408 | Jinghui Qu, Shenzhen, China |
| C*03:198 | D10104742 | KF175409 | Jinghui Qu, Shenzhen, China |
| C*03:199 | D10072405 | KF175410 | Jinghui Qu, Shenzhen, China |
| C*03:200 | D10118127 | KF175411 | Jinghui Qu, Shenzhen, China |
| C*03:201N | D10097621 | KF175412 | Jinghui Qu, Shenzhen, China |
| C*03:202 | D10095948 | KF175413 | Jinghui Qu, Shenzhen, China |
| C*03:203 | D10054540 | KF175414 | Jinghui Qu, Shenzhen, China |
| C*03:204 | D10051069 | KF175416 | Jinghui Qu, Shenzhen, China |
| C*03:205 | D10048588 | KF175417 | Jinghui Qu, Shenzhen, China |
| C*03:206 | D10118603 | KF175418 | Jinghui Qu, Shenzhen, China |
| C*03:207 | D10096862 | KF175419 | Jinghui Qu, Shenzhen, China |
| C*03:208N | D10051727 | KF175420 | Jinghui Qu, Shenzhen, China |
| C*03:209 | D10109671 | KF175423 | Jinghui Qu, Shenzhen, China |
| C*03:210 | D10042999 | KF175425 | Jinghui Qu, Shenzhen, China |
| C*03:211 | D10029002 | KF175428 | Jinghui Qu, Shenzhen, China |
| C*03:212 | D10088501 | KF175429 | Jinghui Qu, Shenzhen, China |
| C*03:213 | D10073288 | KF175430 | Jinghui Qu, Shenzhen, China |
| C*04:01:53 | D10092900 | KF175434 | Jinghui Qu, Shenzhen, China |
| C*04:01:54 | D10048500 | KF175435 | Jinghui Qu, Shenzhen, China |
| C*04:153 | BY01064 | KF266814 | Carolyn K Hurley, Washington DC, USA |
| C*04:154 | D10021513 | KF175433 | Jinghui Qu, Shenzhen, China |
| C*05:95 | D10057122 | KF175438 | Jinghui Qu, Shenzhen, China |
| C*05:96 | D10059824 | KF175437 | Jinghui Qu, Shenzhen, China |
| C*06:02:28 | D10059708 | KF175443 | Jinghui Qu, Shenzhen, China |
| C*06:02:29 | D10109259 | KF175448 | Jinghui Qu, Shenzhen, China |
| C*06:02:30 | D10129772 | KF175449 | Jinghui Qu, Shenzhen, China |
| C*06:103 | P07264 | KF359943 | Faming Zhu, Hangzhou, China |
| C*06:103 | D10064622 | KF175444 | Jinghui Qu, Shenzhen, China |
| C*06:104 | D10059673 | KF175439 | Jinghui Qu, Shenzhen, China |
| C*06:105 | D10082769 | KF175440 | Jinghui Qu, Shenzhen, China |
| C*06:106:01 | D10053009 | KF175441 | Jinghui Qu, Shenzhen, China |
| C*06:106:02 | D10081329 | KF175442 | Jinghui Qu, Shenzhen, China |
| C*06:107 | D10061479 | KF175445 | Jinghui Qu, Shenzhen, China |
| C*06:108 | D10026251 | KF175446 | Jinghui Qu, Shenzhen, China |
| C*06:109 | D10127705 | KF175447 | Jinghui Qu, Shenzhen, China |
| C*06:110 | D10117867 | KF175450 | Jinghui Qu, Shenzhen, China |
| C*06:111 | D10023732 | KF175451 | Jinghui Qu, Shenzhen, China |
| C*07:02:44 | D10064815 | KF175454 | Jinghui Qu, Shenzhen, China |
| C*07:02:45 | D10044257 | KF175457 | Jinghui Qu, Shenzhen, China |
| C*07:02:46 | D10070500 | KF175460 | Jinghui Qu, Shenzhen, China |
| C*07:02:47 | D10128702 | KF175461 | Jinghui Qu, Shenzhen, China |
| C*07:02:48 | D10051528 | KF175462 | Jinghui Qu, Shenzhen, China |
| C*07:02:49 | D10097615 | KF175463 | Jinghui Qu, Shenzhen, China |
| C*07:02:50 | D10064431 | KF175464 | Jinghui Qu, Shenzhen, China |
| C*07:02:51 | D10057081 | KF175465 | Jinghui Qu, Shenzhen, China |
| C*07:316 | 101355 | KF208637 | Chun-Ying Lin, Taipei, Taiwan |
| C*07:317 | BY01062 | KF266816 | Carolyn K Hurley, Washington DC, USA |
| C*07:318 | D10106275 | KF175455 | Jinghui Qu, Shenzhen, China |
| C*07:319 | D10109996 | KF175456 | Jinghui Qu, Shenzhen, China |
| C*07:320 | D10095664 | KF175458 | Jinghui Qu, Shenzhen, China |
| C*07:321 | D10103159 | KF175459 | Jinghui Qu, Shenzhen, China |
| C*07:322 | D10099231 | KF175466 | Jinghui Qu, Shenzhen, China |
| C*07:323 | D10116718 | KF175467 | Jinghui Qu, Shenzhen, China |
| C*07:324 | D10054705 | KF175468 | Jinghui Qu, Shenzhen, China |
| C*08:01:11 | D10067403 | KF175474 | Jinghui Qu, Shenzhen, China |
| C*08:01:12 | D10067254 | KF175475 | Jinghui Qu, Shenzhen, China |
| C*08:79 | D10083171 | KF175469 | Jinghui Qu, Shenzhen, China |
| C*08:80 | D10127194 | KF175470 | Jinghui Qu, Shenzhen, China |
| C*08:81 | D10109077 | KF175471 | Jinghui Qu, Shenzhen, China |
| C*08:82 | D10062302 | KF175473 | Jinghui Qu, Shenzhen, China |
| C*08:83 | D10126572 | KF175476 | Jinghui Qu, Shenzhen, China |
| C*08:84 | D10123391 | KF175477 | Jinghui Qu, Shenzhen, China |
| C*08:85 | D10089657 | KF175478 | Jinghui Qu, Shenzhen, China |
| C*08:86 | D10025458 | KF175479 | Jinghui Qu, Shenzhen, China |
| C*08:87 | D10075175 | KF175480 | Jinghui Qu, Shenzhen, China |
| C*08:88N | D10102176 | KF175481 | Jinghui Qu, Shenzhen, China |
| C*08:89N | D10028151 | KF175482 | Jinghui Qu, Shenzhen, China |
| C*14:53 | JMDP36K0181 | AB829606 | Kenji Tadokoro, Tokyo, Japan |
| C*14:54 | JMDP36K0182 | AB829607 | Kenji Tadokoro, Tokyo, Japan |
| DRB1*04:164 | BY01057 | KF266808 | Carolyn K Hurley, Washington DC, USA |
| DRB1*09:21 | TU11 | AB829527 | Takashi Shiina, Isehara, Japan |
| DRB1*11:148 | BY01058 | KF266807 | Carolyn K Hurley, Washington DC, USA |
| DQB1*03:03:08 | H130270 | KF376099 | Faming Zhu, Hangzhou, China |
| DQB1*03:90N | D10088903 | KF270519 | Jinghui Qu, Shenzhen, China |
| DQB1*06:02:11 | D10047249 | HE804255 | Jinghui Qu, Shenzhen, China |
| DQB1*06:02:12 | D10115720 | HE804260 | Jinghui Qu, Shenzhen, China |
| DQB1*06:03:10 | D10048428 | HE804261 | Jinghui Qu, Shenzhen, China |
| DQB1*06:04:08 | D10116697 | HE804263 | Jinghui Qu, Shenzhen, China |
| DQB1*06:09:04 | D10110619 | HE804265 | Jinghui Qu, Shenzhen, China |
| DQB1*06:106 | D10084766 | HE804256 | Jinghui Qu, Shenzhen, China |
| DQB1*06:107 | D10074295 | HE804257 | Jinghui Qu, Shenzhen, China |
| DQB1*06:108 | D10111798 | HE804254 | Jinghui Qu, Shenzhen, China |
| DQB1*06:109 | D10043019 | HE804259 | Jinghui Qu, Shenzhen, China |
Table 2: Confirmatory Sequences
| Sequence | Cell identification | Accession number | Submitting author |
|---|---|---|---|
| A*23:56 | BY01087 | KF278712 | Carolyn K Hurley, Washington DC, USA |
| A*24:164 | JMDP01K0145 | AB829609 | Kenji Tadokoro, Tokyo, Japan |
| A*24:206 | BY01085 | KF278714 | Carolyn K Hurley, Washington DC, USA |
| A*26:52 | JMDP36K0189 | AB829893 | Kenji Tadokoro, Tokyo, Japan |
| B*15:03:04 | BY01083 | KF266790 | Carolyn K Hurley, Washington DC, USA |
| B*15:241 | BY01070 | KF266803 | Carolyn K Hurley, Washington DC, USA |
| B*35:01:27 | BY01079 | KF266794 | Carolyn K Hurley, Washington DC, USA |
| B*35:11:03 | HG00005995 | KC991059 | Histogenetics, Ossining, USA |
| B*37:35 | BY01080 | KF266793 | Carolyn K Hurley, Washington DC, USA |
| B*40:06:07 | 5761200534 | KF206119 | Faming Zhu, Hangzhou, China |
| B*40:175 | BY01078 | KF266795 | Carolyn K Hurley, Washington DC, USA |
| B*40:86 | JMDP36K0195 | AB829608 | Kenji Tadokoro, Tokyo, Japan |
| B*44:146 | BY01050 | KF040405 | Carolyn K Hurley, Washington DC, USA |
| B*49:23 | BY01077 | KF266796 | Carolyn K Hurley, Washington DC, USA |
| B*51:127 | BY01073 | KF266800 | Carolyn K Hurley, Washington DC, USA |
| C*03:02:10 | D10017137 | KF175407 | Jinghui Qu, Shenzhen, China |
| C*03:86 | BY01066 | KF266812 | Carolyn K Hurley, Washington DC, USA |
| C*06:03:02 | BY01069 | KF266809 | Carolyn K Hurley, Washington DC, USA |
| C*06:101 | BY01063 | KF266815 | Carolyn K Hurley, Washington DC, USA |
| C*08:01:07 | BY01065 | KF266813 | Carolyn K Hurley, Washington DC, USA |
| C*08:01:10 | D10046648 | KF175472 | Jinghui Qu, Shenzhen, China |
| C*08:02:04 | BY01068 | KF266810 | Carolyn K Hurley, Washington DC, USA |
| DRB1*04:154 | BY01059 | KF266806 | Carolyn K Hurley, Washington DC, USA |
| DRB1*14:94 | BY01055 | KF266805 | Carolyn K Hurley, Washington DC, USA |
| DQB1*06:84 | D10080597 | HE804258 | Jinghui Qu, Shenzhen, China |
Table 3: Recently Published Sequences
| Sequence | References |
|---|---|
| A*01:128 | (3) |
| A*02:356N | (4) |
| A*02:425 | (5) |
| A*24:183N | (4) |
| A*24:215 | (6) |
| A*24:223 | (7) |
| A*33:44 | (8) |
| B*08:57 | (9) |
| B*15:139 | (10) |
| B*27:07:02 | (11) |
| B*35:206 | (11) |
| B*35:226 | (12) |
| B*40:06:06 | (13) |
| B*40:221 | (14) |
| B*40:223 | (12) |
| B*58:01:12 | (15) |
| B*58:43 | (16) |
| C*03:190 | (16) |
| C*05:88 | (3) |
| DRB1*04:05:14 | (17) |
| DRB1*04:05:15 | (18) |
| DRB1*09:01:07 | (19) |
| DRB1*13:26:02 | (20) |
| DQB1*04:10 | (21) |
| DQB1*05:19 | (22) |
| DQB1*06:48 | (23) |
| DPA1*01:12 | (16) |
| MICB*005:07 | (24) |
| MICB*005:08 | (24) |
| MICB*027 | (24) |
| MICB*028 | (24) |
| MICB*029 | (24) |
References:
- Marsh SGE, Albert ED, Bodmer WF, et al. Nomenclature for Factors of the HLA system, 2010. Tissue Antigens. 2010: 75:291-455.
- Robinson J, Malik A, Parham P, Bodmer JG, Marsh SGE. IMGT/HLA database--a sequence database for the human major histocompatibility complex. Tissue Antigens. 2000: 55:280-7.
- Sanchez-Gordo F, Balas A, Gomez-Zumaquero JL, Prat I, Vicario JL. Description of two new HLA alleles, A*01:128 and C*05:88, identified in Spanish individuals. Tissue Antigens. 2013: 82: 134-5.
- Shimizu M, Nakajima F, Okazaki H, Satake M, Tadokoro K. Two novel null HLA-A alleles with identical exon 4 nonsense mutations: HLA-A*24:183N and A*02:356N. Tissue Antigens. 2013: 82: 136-7.
- Huo MR, Yu Y, Xi B, Li D. Identification of a novel HLA-A*02 allele, HLA-A*02:425, by sequence-based typing. Tissue Antigens. 2013: 82: 135-6.
- Plaisier L, Geerts N, van Ginkel W, van der Weerd R, Spierings E. The novel HLA-A*24:215 allele, identified by sequencing-based typing of a stem cell transplant patient and the sibling donor. Tissue Antigens. 2013: 82: 138-9.
- Wang D, Xu Y, Zou H, Deng Z. A novel HLA-A allele detected by sequence-based typing: HLA-A*24:223. Tissue Antigens. 2013: 82: 139-40.
- Ye X, Ding H, Huang YF, Tan Y, Xia W, Bei C. HLA-A*33:44: a new allele identified by sequence-based typing in a Chinese blood donor. Tissue Antigens. 2013: 82: 140-2.
- Zhang X, Li XF, Lin FQ, Zhang KL, Li JP. Detection of the new HLA-B*08:57 allele in a voluntary platelet donor. Tissue Antigens. 2013: 82: 144-5.
- Zhang WD, Wang LJ, Zhang KY, Shan XY, Jia YJ. Characterization of a novel allele, HLA-B*15:139. Tissue Antigens. 2013: 82: 145-6.
- Frison S, Longhi E, Mantovani M, et al. Two novel alleles at HLA-B locus identified in two volunteer bone marrow donors by sequence-based typing. International Journal of Immunogenetics. 2013: 40: 328-30.
- Luo J, Yang N, Cao LQ, Xu ZX. Two new alleles, HLA-B*35:226 and HLA-B*40:233 identified by sequence-based typing in Chinese individuals. Tissue Antigens. 2013: 82: 147-8.
- Zou HY, Jin SZ, Li Z. HLA-B*40:06:06, a novel allele, which has arisen by silent mutation in codon 45. Tissue Antigens. 2013: 82: 142-4.
- Yang KL, Lee SK, Lin PY. Identification of the novel HLA-B*40:221 allele in a Taiwanese hematopoietic stem cell donor using a sequence-based typing method. International Journal of Immunogenetics. 2013: 40: 320-1.
- Yang KL, Lee SK, Kao RH, Lin CL, Lin PY. A novel HLA-B allele, B*58:01:12, detected in a Taiwanese volunteer bone marrow stem cell donor using sequence-based typing method. International Journal of Immunogenetics. 2013: 40: 324-5.
- Nykoluk M, Bailey RC, Moses S, Plummer FA, Luo M. Identification of three novel human leukocyte antigen alleles, HLA-B*58:43, HLA-C*03:190, and HLA-DPA1*01:12, in an East African cohort. Tissue Antigens. 2013: 82: 131-3.
- Yang KL, Lee SK, Kao RH, Lin CL, Lin PY. Discovery of the novel HLA-DRB1*04:05:14 allele in a Taiwanese unrelated haematopoietic stem cell donor by a sequence-based typing method. International Journal of Immunogenetics. 2013: 40: 326-7.
- Yang KL, Lee SK, Lin PY. Discovery of a novel HLA-DRB1*04:05 variant, DRB1*04:05:15, in a Taiwanese and the probable HLA haplotype in association with DRB1*04:05:15. International Journal of Immunogenetics. 2013: 40: 396-7.
- Li XF, Zhang X, Zhang KL, Liu XZ, Li JP. Sequencing of a novel HLA-DRB1*09:01:07 allele in a Chinese individual. Tissue Antigens. 2013: 82: 151-2.
- Zheng JW, Yang JH, Xie JH, et al. Characterization of the novel HLA-DRB1*13:26:02 allele. Tissue Antigens. 2013: 82: 152-3.
- Oka S, Furukawa H, Kashiwase K, Tsuchiya N, Tohma S. A novel HLA-DQB1*04 allele, DQB1*04:10, identified in a Japanese individual. Tissue Antigens. 2013: 82: 148-9.
- Hommel-Berrey GA, House KD, Backfish KA, Arnoff TS, Lobashevsky A. Identification of a novel HLA-DQB1 allele, DQB1*05:19, in an African American family by sequence-based typing. Tissue Antigens. 2013: 82: 150-1.
- Foster L, Tate D, Poulton K. Characterization of the novel HLA-DQB1*06:48 allele by group-specific sequencing. International Journal of Immunogenetics. 2013: 40: 322-3.
- Ying Y, He Y, Tao S, et al. Distribution of MICB diversity in the Zhejiang Han population: PCR sequence-based typing for exons 2-6 and identification of five novel MICB alleles. Immunogenetics. 2013: 65: 485-92.
