Nomenclature for Factors of the HLA System

September 2012

Compiled by Steven G. E. Marsh for the WHO Nomenclature Committee for Factors of the HLA System.

Published In:

The following sequences have been submitted to the Nomenclature Committee since the August 2012 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. The sequences A*02:01:82, A*24:211 and C*08:64 were shown to have been named in error and be identical to A*02:01:84, A*24:135:02 and C*08:33:02 respectively. The names A*02:01:82, A*24:211 and C*08:64 have been abandoned.

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*01:01:50HG00004531JQ964641Histogenetics, Ossining, USA
A*01:01:510231200743JX629459Wei Mao, Chongqing, China
A*01:83:02HG00004534JQ867313Histogenetics, Ossining, USA
A*01:120HG00004530JQ935214Histogenetics, Ossining, USA
A*01:121HG00004532JX458873Histogenetics, Ossining, USA
A*01:122HG00004533JX458872Histogenetics, Ossining, USA
A*02:01:85HG00004545JQ911862Histogenetics, Ossining, USA
A*02:01:86HG00004546JQ911858Histogenetics, Ossining, USA
A*02:387HG00004539JQ824426Histogenetics, Ossining, USA
A*02:388HG00004540JX390630Histogenetics, Ossining, USA
A*02:389HG00004541JX441341Histogenetics, Ossining, USA
A*02:390HG00004542JX441337Histogenetics, Ossining, USA
A*02:391HG00004543JQ997114Histogenetics, Ossining, USA
A*02:392HG00004544JX267233Histogenetics, Ossining, USA
A*03:01:38HG00004535JX267234Histogenetics, Ossining, USA
A*03:160HG00004536JX390625Histogenetics, Ossining, USA
A*03:161NHG00004538JQ935204Histogenetics, Ossining, USA
A*11:131HG00004504JX290325Histogenetics, Ossining, USA
A*11:132HG00004510JQ935211Histogenetics, Ossining, USA
A*23:01:11HG00004520JX412248Histogenetics, Ossining, USA
A*23:55HG00004522JX441340Histogenetics, Ossining, USA
A*24:02:62HG00004527JX290326Histogenetics, Ossining, USA
A*24:02:63HG00004528JQ911861Histogenetics, Ossining, USA
A*24:10:02HG00004519JX458870Histogenetics, Ossining, USA
A*24:135:02HG00002960JN124111Histogenetics, Ossining, USA
A*24:215522-42JX564252Talitha de Hoop, Utrecht, The Netherlands
A*24:216HG00004517JQ935207Histogenetics, Ossining, USA
A*24:217HG00004518JQ997111Histogenetics, Ossining, USA
A*24:218HG00004523JX412247Histogenetics, Ossining, USA
A*24:219HG00004524JX027032Histogenetics, Ossining, USA
A*24:220HG00004525,
HG00004526
JX228000,
JX228001
Histogenetics, Ossining, USA
A*24:221HG00004529JQ997108Histogenetics, Ossining, USA
A*24:222NBY00942JX674058Carolyn K Hurley, Washington DC, USA
A*25:22HG00004506JX441334Histogenetics, Ossining, USA
A*26:01:27HG00004503JX227997Histogenetics, Ossining, USA
A*26:795751100703JX524131Faming Zhu, Hangzhou, China
A*26:80HG00004496JQ824428Histogenetics, Ossining, USA
A*29:46553-43JX564250Talitha de Hoop, Utrecht, The Netherlands
A*29:471-10265JX679217Carmen Alaez, Mexico, Mexico
A*30:02:09HG00004514JX290330Histogenetics, Ossining, USA
A*30:02:10HG00004515JQ911859Histogenetics, Ossining, USA
A*30:67HG00004516JX227996Histogenetics, Ossining, USA
A*30:6822904493HE983832Thibaut Gervais, Brussels, Belgium
A*30:69BY00934JX674066Carolyn K Hurley, Washington DC, USA
A*30:70NBY00947JX674053Carolyn K Hurley, Washington DC, USA
A*31:68HG00004497JQ867305Histogenetics, Ossining, USA
A*31:69HG00004498JX390628Histogenetics, Ossining, USA
A*31:70HG00004499JQ997112Histogenetics, Ossining, USA
A*32:33:02BY00945JX674055Carolyn K Hurley, Washington DC, USA
A*32:55HG00004537JQ911860Histogenetics, Ossining, USA
A*33:03:1120110278JX524132Faming Zhu, Hangzhou, China
A*33:03:12BY00944JX674056Carolyn K Hurley, Washington DC, USA
A*33:03:13SZ-151JX846910Hong-Yan Zou, Shenzhen, China
A*33:63HG00004500JX267235Histogenetics, Ossining, USA
A*33:64HG00004501JX441339Histogenetics, Ossining, USA
A*34:10NHG00004508JX267230Histogenetics, Ossining, USA
A*66:19HG00004507JX290329Histogenetics, Ossining, USA
A*68:01:17HG00004512JX441336Histogenetics, Ossining, USA
A*68:13:02HG00004513JX412245Histogenetics, Ossining, USA
A*68:95HG00004511JQ911864Histogenetics, Ossining, USA
A*68:96CA6690HE984353Xiang Liu, Beijing, China
A*80:01:01:02CTM-4687797JX826590Antonio Balas, Madrid, Spain
B*07:02:36HG00004583JQ867324Histogenetics, Ossining, USA
B*07:02:37HG00004585JQ935218Histogenetics, Ossining, USA
B*07:176HG00004582JX228021Histogenetics, Ossining, USA
B*08:01:19HG00004563JQ911878Histogenetics, Ossining, USA
B*08:01:20FRF1658HE974341Sylvie Tourne, Strasbourg, France
B*08:53:02HG00004565JX228004Histogenetics, Ossining, USA
B*08:102HG00004564JQ824440Histogenetics, Ossining, USA
B*08:103HG00004566JX390631Histogenetics, Ossining, USA
B*08:104HG00004567JX441351Histogenetics, Ossining, USA
B*08:105BY00940JX674060Carolyn K Hurley, Washington DC, USA
B*13:62HG00004569JX027042Histogenetics, Ossining, USA
B*13:63NR28817HE984577Edward KL Yang, Hualien, Taiwan
B*15:01:31HG00004576JX412252Histogenetics, Ossining, USA
B*15:263KJ, 6544631JX470440Oh-Joong Kwon, Seoul, South Korea
B*15:263CB11-468JX646672Ji Young Huh, Sungnam-si, South Korea
B*15:264BY00939JX674061Carolyn K Hurley, Washington DC, USA
B*18:01:18HG00004559JX441350Histogenetics, Ossining, USA
B*18:83HG00004558JX458861Histogenetics, Ossining, USA
B*27:05:20HG00004556JX110626Histogenetics, Ossining, USA
B*27:99HG00004557JQ824436Histogenetics, Ossining, USA
B*35:01:36HG00004574JQ997134Histogenetics, Ossining, USA
B*35:03:12HG00004575JX441343Histogenetics, Ossining, USA
B*35:61:0210120Y33JX674062Carolyn K Hurley, Washington DC, USA
B*35:219BY00941JX674059Carolyn K Hurley, Washington DC, USA
B*35:220BS1741HE984354Xiang Liu, Beijing, China
B*37:37BY00935JX674065Carolyn K Hurley, Washington DC, USA
B*39:01:15RH125JX284120Yann-Jinn Lee, Taipei County, Taiwan
B*40:06:06SZ-150JX493014Hong-Yan Zou, Shenzhen, China
B*40:224HG00004553JX267236Histogenetics, Ossining, USA
B*40:225HG00004554JQ935216Histogenetics, Ossining, USA
B*40:226HG00004555JQ997129Histogenetics, Ossining, USA
B*41:23HG00004548JX228023Histogenetics, Ossining, USA
B*41:24HG00004549JX458864Histogenetics, Ossining, USA
B*41:25HG00004551JX458865Histogenetics, Ossining, USA
B*44:02:27557-46JX564251Talitha de Hoop, Utrecht, The Netherlands
B*44:59:02HG00004571JX267239Histogenetics, Ossining, USA
B*44:162HG00004570JX267238Histogenetics, Ossining, USA
B*44:163HG00004572JX228020Histogenetics, Ossining, USA
B*44:164HG00004581JQ867326Histogenetics, Ossining, USA
B*44:165HG00004586JX412251Histogenetics, Ossining, USA
B*46:38CA5287HE984356Xiang Liu, Beijing, China
B*49:24E000911002985JX645189Antonio Balas, Madrid, Spain
B*49:25HG00004568JX267242Histogenetics, Ossining, USA
B*50:19HG00004550JX441349Histogenetics, Ossining, USA
B*51:01:38HG00004578JX228005Histogenetics, Ossining, USA
B*51:145HG00004547JX228018Histogenetics, Ossining, USA
B*51:146HG00004579JX267241Histogenetics, Ossining, USA
B*51:147HG00004580JQ824437Histogenetics, Ossining, USA
B*51:148BY00937JX674063Carolyn K Hurley, Washington DC, USA
B*52:01:16HG00004573JX290332Histogenetics, Ossining, USA
B*55:01:10HG00004577JX228003Histogenetics, Ossining, USA
C*02:02:25HG00004666JQ824581Histogenetics, Ossining, USA
C*02:65HG00004667JQ911924Histogenetics, Ossining, USA
C*02:66HG00004668JQ824666Histogenetics, Ossining, USA
C*03:04:32HG00004678JQ911907Histogenetics, Ossining, USA
C*03:172HG00004677JQ824488Histogenetics, Ossining, USA
C*03:173HG00004679JQ824687Histogenetics, Ossining, USA
C*03:174HG00004680JQ824705Histogenetics, Ossining, USA
C*03:175HG00004682JQ824700Histogenetics, Ossining, USA
C*05:01:24HG00004639JQ824567Histogenetics, Ossining, USA
C*05:01:25HG00004641JQ824466Histogenetics, Ossining, USA
C*05:04:02HG00004645JQ824595Histogenetics, Ossining, USA
C*05:29:02HG00004660JX458890Histogenetics, Ossining, USA
C*05:82HG00004642JQ824529Histogenetics, Ossining, USA
C*05:83HG00004643JQ824514Histogenetics, Ossining, USA
C*05:84HG00004644JQ824704Histogenetics, Ossining, USA
C*05:85HG00004646JQ824545Histogenetics, Ossining, USA
C*06:02:22HG00004598JX458886Histogenetics, Ossining, USA
C*06:02:23HG00004599JQ824542Histogenetics, Ossining, USA
C*06:02:24HG00004603JX228031Histogenetics, Ossining, USA
C*06:88HG00004593JQ824644Histogenetics, Ossining, USA
C*06:89HG00004595JQ824609Histogenetics, Ossining, USA
C*06:90HG00004596JQ824536Histogenetics, Ossining, USA
C*06:91HG00004597JQ824523Histogenetics, Ossining, USA
C*06:92HG00004600JX228052Histogenetics, Ossining, USA
C*06:93HG00004602JQ824608Histogenetics, Ossining, USA
C*07:01:29HG00004611JX458878Histogenetics, Ossining, USA
C*07:01:30HG00004613,
HG00004614
JX267243,
JQ824487
Histogenetics, Ossining, USA
C*07:01:31HG00004616JQ824593Histogenetics, Ossining, USA
C*07:02:37HG00004630JQ824600Histogenetics, Ossining, USA
C*07:02:38HG00004631JQ824443Histogenetics, Ossining, USA
C*07:02:39HG00004636JQ824528Histogenetics, Ossining, USA
C*07:17:02HG00004629JQ824552Histogenetics, Ossining, USA
C*07:96:02HG00004675JQ824588Histogenetics, Ossining, USA
C*07:271HG00004588JQ824576Histogenetics, Ossining, USA
C*07:272HG00004589JX458885Histogenetics, Ossining, USA
C*07:273HG00004591JQ824662Histogenetics, Ossining, USA
C*07:274HG00004592JX228032Histogenetics, Ossining, USA
C*07:275HG00004604JX458889Histogenetics, Ossining, USA
C*07:276HG00004605JQ911912Histogenetics, Ossining, USA
C*07:277HG00004606JQ824622Histogenetics, Ossining, USA
C*07:278HG00004607JQ824478Histogenetics, Ossining, USA
C*07:279HG00004608,
HG00004609
JX458891,
JQ964668
Histogenetics, Ossining, USA
C*07:280HG00004610JQ824500Histogenetics, Ossining, USA
C*07:281HG00004612JQ824587Histogenetics, Ossining, USA
C*07:282HG00004615JQ824568Histogenetics, Ossining, USA
C*07:283HG00004617JQ824623Histogenetics, Ossining, USA
C*07:284HG00004618JQ911910Histogenetics, Ossining, USA
C*07:285HG00004619,
HG00004620,
HG00004621,
HG00004622,
HG00004623
JQ824446,
JQ824556,
JQ824557,
JQ867345,
JQ964671
Histogenetics, Ossining, USA
C*07:286HG00004626JX458895Histogenetics, Ossining, USA
C*07:287HG00004627JX228030Histogenetics, Ossining, USA
C*07:288HG00004628JQ824611Histogenetics, Ossining, USA
C*07:289HG00004632JQ911917Histogenetics, Ossining, USA
C*07:290HG00004633JQ824598Histogenetics, Ossining, USA
C*07:291HG00004634JQ824589Histogenetics, Ossining, USA
C*07:292HG00004635JQ964659Histogenetics, Ossining, USA
C*08:02:08HG00004664,
HG00004665
JQ824448,
JQ824507
Histogenetics, Ossining, USA
C*08:04:03HG00004649JQ824471Histogenetics, Ossining, USA
C*08:33:03HG00003938JQ007713Histogenetics, Ossining, USA
C*08:67HG00004637JX458876Histogenetics, Ossining, USA
C*08:68HG00004638JX458883Histogenetics, Ossining, USA
C*08:69HG00004640JQ824517Histogenetics, Ossining, USA
C*08:70HG00004647JQ824546Histogenetics, Ossining, USA
C*08:71HG00004648JQ824467Histogenetics, Ossining, USA
C*08:72HG00004651JQ867331Histogenetics, Ossining, USA
C*12:03:24HG00004652JQ911925Histogenetics, Ossining, USA
C*12:03:25HG00004663JQ824456Histogenetics, Ossining, USA
C*12:86HG00004655JX228051Histogenetics, Ossining, USA
C*12:87HG00004656JX458893Histogenetics, Ossining, USA
C*12:88HG00004657JQ824615Histogenetics, Ossining, USA
C*12:89HG00004658JX458894Histogenetics, Ossining, USA
C*12:90HG00004659JQ824640Histogenetics, Ossining, USA
C*12:91HG00004662JQ824620Histogenetics, Ossining, USA
C*15:02:11HG00004670JQ824457Histogenetics, Ossining, USA
C*15:02:12HG00004671JQ824532Histogenetics, Ossining, USA
C*15:02:13HG00004672JQ824676Histogenetics, Ossining, USA
C*15:02:14HG00004673JQ824672Histogenetics, Ossining, USA
C*15:02:15HG00004676JX458882Histogenetics, Ossining, USA
C*15:63HG00004594JX458887Histogenetics, Ossining, USA
C*15:64HG00004674JQ824601Histogenetics, Ossining, USA
C*16:01:11HG00004654JQ911897Histogenetics, Ossining, USA
C*17:15HG00004661JX228050Histogenetics, Ossining, USA
C*18:06HG00004590JX267246Histogenetics, Ossining, USA
DRB1*01:01:23HG00004780JQ824710Histogenetics, Ossining, USA
DRB1*01:5010-72-1133HE983354Wendy Swelsen, Amsterdam, The Netherlands
DRB1*01:51HG00004781JQ824714Histogenetics, Ossining, USA
DRB1*01:52NHG00004782JQ824707Histogenetics, Ossining, USA
DRB1*01:53HG00004783JQ824711Histogenetics, Ossining, USA
DRB1*03:86HG00004779JQ824804Histogenetics, Ossining, USA
DRB1*04:01:10DKMS-LSL-DRB1-11HE993875DKMS Life Science Lab, Dresden, Germany
DRB1*04:128DKMS-LSL-DRB1-7HE993871DKMS Life Science Lab, Dresden, Germany
DRB1*04:129DKMS-LSL-DRB1-9HE993873DKMS Life Science Lab, Dresden, Germany
DRB1*04:130DKMS-LSL-DRB1-10HE993874DKMS Life Science Lab, Dresden, Germany
DRB1*04:131DKMS-LSL-DRB1-12HE993876DKMS Life Science Lab, Dresden, Germany
DRB1*04:132DKMS-LSL-DRB1-13HE993877DKMS Life Science Lab, Dresden, Germany
DRB1*04:133DKMS-LSL-DRB1-14HE993878DKMS Life Science Lab, Dresden, Germany
DRB1*11:134HG00004778JQ824779Histogenetics, Ossining, USA
DRB1*14:05:04D10112187HE654761Jinghui Qu, Shenzhen, China
DRB1*14:130D10058865HE654762Jinghui Qu, Shenzhen, China
DRB1*14:131D10106977HE654764Jinghui Qu, Shenzhen, China
DRB1*15:01:21D10113968HE654767Jinghui Qu, Shenzhen, China
DRB1*15:37:02D10128181HE654768Jinghui Qu, Shenzhen, China
DRB1*15:81D10026958HE654766Jinghui Qu, Shenzhen, China
DQB1*02:01:0611-8205JX471069Dimitri Monos, Philadelphia, USA
DQB1*05:17C149354JX494227Robert Endres, Tempe, USA
DQB1*05:1809-72-3127HE983355Wendy Swelsen, Amsterdam, The Netherlands
DPA1*02:02:05SZ-QJHJX549436Yunping Xu, Shenzhen, China
DPB1*02:01:08DKMS-LSL-DPB1-15HE995398DKMS Life Science Lab, Dresden, Germany
DPB1*141:01DKMS-LSL-DPB1-16HE995399DKMS Life Science Lab, Dresden, Germany

Table 2: Confirmatory Sequences


Sequence Cell identification Accession number Submitting author
A*01:64BY00949JX6704051Carolyn K Hurley, Washington DC, USA
A*02:01:84BY00930JX423801Carolyn K Hurley, Washington DC, USA
A*02:28338769HE983615Edward KL Yang, Hualien, Taiwan
A*11:01:19HG00004505JQ911863Histogenetics, Ossining, USA
A*11:70HG00004509JQ824424Histogenetics, Ossining, USA
A*23:01:03HG00004521JQ911857Histogenetics, Ossining, USA
A*24:02:0523122864HE983831Thibaut Gervais, Brussels, Belgium
A*26:01:18BY00948JX674052Carolyn K Hurley, Washington DC, USA
A*32:44BY00943JX674057Carolyn K Hurley, Washington DC, USA
A*33:42BY00946JX674054Carolyn K Hurley, Washington DC, USA
B*07:175HG00004584JQ867325Histogenetics, Ossining, USA
B*18:54HG00004560JQ964646Histogenetics, Ossining, USA
B*18:68HG00004552JQ911881Histogenetics, Ossining, USA
B*18:79HG00004561,
HG00004562
JX267240,
JX228022
Histogenetics, Ossining, USA
B*40:01:25BX1978HE984355Xiang Liu, Beijing, China
C*01:54HG00004587JQ997151Histogenetics, Ossining, USA
C*01:6709-72-3827HE983356Wendy Swelsen, Amsterdam, The Netherlands
C*03:10D223480HE983616Edward KL Yang, Hualien, Taiwan
C*03:85HG00004681JQ911900Histogenetics, Ossining, USA
C*07:04:06HG00004601JQ867332Histogenetics, Ossining, USA
C*07:149HG00004625JQ911914Histogenetics, Ossining, USA
C*07:171HG00004624JQ824544Histogenetics, Ossining, USA
C*08:56HG00004650JQ824484Histogenetics, Ossining, USA
C*15:06:03HG00004669JQ867346Histogenetics, Ossining, USA
DQA1*01:07215443HE980019Klaus Witter, München, Germany
DQA1*03:02215443HE980020Klaus Witter, München, Germany

Table 3: Recently Published Sequences


Sequence References
A*24:132N(3)
A*24:96(4)
A*31:22(5)
A*33:22(6)
B*13:19(7)
B*35:189(8)
B*40:186(9)
C*04:107(10)
C*07:244(11)
G*01:18(12)
MICA*066(13)
DQB1*06:49(14)
DQB1*06:51(15)
DPB1*136:01(16)

References:

  1. Marsh SGE, Albert ED, Bodmer WF, et al. Nomenclature for Factors of the HLA system, 2010. Tissue Antigens. 2010 75:291-455.
  2. 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.
  3. Zou HY, Jin SZ, Li Z. A HLA-A null allele (A*24:132N) with a stop codon in exon 3 generated by a point mutation. Tissue Antigens. 2012: 80: 464-5.
  4. Wang L, Zhixin Z, Xiaoyan S, Zhao B, Gong Z. Identification of the novel HLA-A allele, HLA-A*24:96, in a Chinese individual. Tissue Antigens. 2012: 80: 462-3.
  5. Zhang WW, Li XF, Zhang X, Li JP, Li ZQ. A novel HLA-A31 allele, A*31:22, was identified by sequence-based typing. Tissue Antigens. 2012: 80: 380-1.
  6. Xie JH, Yan LN, Li SY, An SP, Yuan YH. Identification of the novel allele HLA-A*33:22 by sequence-based typing in a Chinese individual. Tissue Antigens. 2012: 80: 381-3.
  7. Wang L, Zhixin Z, Xiaoyan S, Dongmei W, Li D. Identification of a novel HLA-B allele, HLA-B*13:19, in a Chinese individual. Tissue Antigens. 2012: 80: 466-7.
  8. Nie XM, Zhang Y, Zhuang YL, Song YH, Zhu CF. A novel HLA-B*35 allele, B*35:189 with a new motif GGG at codon 35, identified by sequence-based typing in a Chinese potential donor. Tissue Antigens. 2012: 80: 467-9.
  9. Kim SY, Shin KH, Yoon CE, Kwon OJ, Kim HH. A new HLA-B*40:186 allele identified by sequence-based typing in a Korean individual. Tissue Antigens. 2012: 80: 383-4.
  10. Jin SZ, Zou HY. Identification of a novel HLA-C*04 allele, HLA-C*04:107, in a Chinese individual. Tissue Antigens. 2012: 80: 469-71.
  11. Cho D, Jang MJ, Yoon CE, Kwon OJ, Shin MG. A new HLA-C*07:244 allele identified by sequence-based typing in a Korean individual. Tissue Antigens. 2012: 80: 384-5.
  12. Roger J, Faucher MC, Roger M. Identification of a new HLA-G allele, HLA-G*01:18, in a Canadian Caucasian individual. Tissue Antigens. 2012: 80: 472-3.
  13. Gao SQ, Wang XT, Xu YP, Deng ZH. MICA*066, a novel major histocompatibility complex class I-related chain A allele found by high-resolution HLA-matching tests for hematopoietic stem cell transplantation. Tissue Antigens. 2012: 80: 473-5.
  14. Todorova K, Diederich G, Ebell W, Kuhl JS, Schonemann C, Schulze H. Detection of a novel HLA-DQB1 allele, designated DQB1*06:49. Tissue Antigens. 2012: 80: 387-8.
  15. Oka S, Furukawa H, Kashiwase K, Tsuchiya N, Tohma S. Identification of a novel HLA allele, HLA-DQB1*06:51, in a Japanese rheumatoid arthritis patient. Tissue Antigens. 2012: 80: 386-7.
  16. Abad ML, Balas A, Schmidt AH, Rodriguez-Calvo MI, Vicario JL, Eiras A. A novel allele HLA-DPB1*136:01 identified in a German hematopoietic stem cell volunteer donor of Turkish descent. Tissue Antigens. 2012: 80: 471-2.