Citing IMGT databases: Manso T. et al. IMGT® databases, related tools and web resources through three main axes of research and development. Nucleic Acids Res. 2022 Jan 7;50(D1):D1262-D1272. doi: 10.1093/nar/gkab1136 PMID: 34875068 Free PMC article.
Program version: v. 

Add the possibility to obtain a gene table per strain (for mouse now and for other species later) and allotypes/isotypes for human.
September 25th, 2024.

Bibliographical references in alphabetic order, small design changes and addition of "NL" for non-localized gene.
June 14th, 2024.

Addition of 'Score for IMGT allele confirmation' as well as NCBI TPA accession numbers.
September 20th, 2023.

Implementation of the dynamic gene table.

Gene table legend:

"+" or "-" indicates if the gene sequences have been found (+) or not been found (-) rearranged (R), transcribed (T) and/or translated into protein (Pr). Arbitrarily that information is shown on the first line of each gene, when the data have been confirmed by several studies.

Functionality is shown in parentheses, (F) and (P), when the accession number refers to rearranged genomic DNA or cDNA and the corresponding germline gene has not yet been isolated.

IMGT allele confirmation: a scoring system is employed to signify the frequency of occurrences of a specific allele within sequence data. A single star () indicates that no instances of the allele have been identified in the literature. Two stars () indicate the discovery of one literature sequence containing the allele. If more than one literature sequence is found to contain the allele, it is designated with three stars (). In the Excel file, the stars are represented by the plus symbol (+).

Click on:
  • IMGT gene name to get the corresponding IMGT/GENE-DB entry (link).
  • IMGT allele name to see the corresponding Alignments of alleles (link).
  • Accession number to get the corresponding IMGT/LIGM-DB entry (link).
  • MAP: mapped sequences. Click to access GENE-DB «LOCALIZATION IN GENOME ASSEMBLIES» (link).
  • [number] to access the corresponding IMGT reference (popover).
  • (number) to see the corresponding IMGT note (popover).
Options:
  • You can show/hide columns (), download data () or put the table in fullscreen () using buttons.
See also (IMGT Scientific chart):
Select a species and a IMGT group to get the gene table:
Only IMGT available species/group are shown in the drop-down list.
The gene table can take several seconds to appear, please be patient.
Gene table of human (Homo sapiens) IGHD IMGT group:
IMGT sub­groupIMGT gene nameIMGT allele nameScore for
IMGT allele
confirmation
FctChromosomosal
localization
R T PrIMGT/LIGM-DB reference sequencesIMGT/LIGM-DB sequences from the literature
Clone namesAccession
numbers
Positions
in the sequence
(D-GENE-UNIT)
or D-REGION (*)
Secondary
accession
numbers
Clone namesAccession
numbers
Positions
in the sequence
(D-GENE-UNIT)
or D-REGION (*)
IGHD1IGHD1-1 IGHD1-1*01 F 14q32.33
RTPr
++
D1-1 X97051 [1,5,13,15,18] MAP 33686-33758 AB019441 [12] 104136-104208
ABC13_11_000049738900_M6 AC279151 [17] (2) complement(3767-3839)
BK063799 [11] 960299-960371
BK063800 [11] 970879-970951
BK063801 [11] 926081-926153
BK063802 [11] 904136-904208
BK068298 [11] 993137-993209
BK068299 [11] 974274-974346
IMGT000054 [17] (2) 35741-35813
IMGT000176 [6-8] 467722-467794
IMGT000177 [9,10] 1071017-1071089
IMGT000207 740748-740820
JADDFX010000192 [19] complement(139091-139163)
IGHD1IGHD1-7 IGHD1-7*01 F 14q32.33
RTPr
++
DM1 X13972 [5] MAP (3) 5238-5310 (4) AB019441 [12] 113731-113803
BK063799 [11] 969894-969966
BK063800 [11] 980473-980545
BK063802 [11] 913731-913803
BK068298 [11] 1002731-1002803
BK068299 [11] 983856-983928
IMGT000177 [9,10] 1080603-1080675
IMGT000207 750343-750415
JADDFX010000192 [19] complement(129497-129569)
M37277 [5] (3) 5266-5282 *
X97051 [1,5,13,15,18] 43272-43344
IGHD1IGHD1-14 IGHD1-14*01 ORF (5) 14q32.33
RTPr
++
DM2 X13972 [5] MAP (3,6) 14490-14562 (4) AB019441 [12] 123001-123073
BK063799 [11] 979163-979235
BK063800 [11] 989742-989814
BK063801 [11] 935326-935398
BK063802 [11] 923001-923073
BK068298 [11] 1011998-1012070
BK068299 [11] 993125-993197
IMGT000207 759612-759684
JADDFX010000192 [19] complement(120351-120423)
M37277 [5] (3,6) 14518-14534 *
X97051 [1,5,13,15,18] 52540-52612
IGHD1IGHD1-20 IGHD1-20*01 F 14q32.33
RTPr
++
D1-20 X97051 [1,5,13,15,18] MAP 61987-62059 AB019441 [12] 132446-132518
BK063799 [11] 988610-988682
BK063800 [11] 999195-999267
BK063801 [11] 944773-944845
BK063802 [11] 932448-932520
BK068298 [11] 1021451-1021523
BK068299 [11] 1002572-1002644
IMGT000207 769059-769131
JADDFX010000192 [19] complement(110898-110970)
IGHD1IGHD1-26 IGHD1-26*01 F 14q32.33
RTPr
++
D1-26 X97051 [1,5,13,15,18] MAP 72141-72216 AB019441 [12] 142598-142673
BK063799 [11] 998764-998839
BK063800 [11] 1009268-1009343
BK063801 [11] 954927-955002
BK063802 [11] 942602-942677
BK068298 [11] 1031523-1031598
BK068299 [11] 1012645-1012720
JADDFX010000192 [19] complement(100980-101055)
IGHD2IGHD2-2 IGHD2-2*01 F 14q32.33
RTPr
++
D4 J00232 [18] MAP 1-87 ABC13_11_000049738900_M6 AC279151 [17] (2) complement(1091-1177)
BK063799 [11] 962961-963047
BK063800 [11] 973541-973627
BK063801 [11] 928742-928828
BK063802 [11] 906798-906884
BK068298 [11] 995799-995885
BK068299 [11] 976924-977010
ABC13_11_000049738900_M6 IMGT000054 [17] (2) 38403-38489
IMGT000207 743410-743496
JADDFX010000192 [19] complement(136415-136501)
IGHD2IGHD2-2 IGHD2-2*02 F 14q32.33
RTPr
++
D2-2 X97051 [1,5,13,15,18] 36339-36425 AB019441 [12] 106798-106884
IMGT000177 [9,10] 1073670-1073756
IGHD2IGHD2-2 IGHD2-2*03 F 14q32.33
RTPr
++
D4 M35648 [20] 42-128 M35649 [20] 70-100 *
IGHD2IGHD2-8 IGHD2-8*01 F 14q32.33
RTPr
++
DLR1 X13972 [5] MAP (3) 7921-8007 (4) AB019441 [12] 116414-116500
BK063799 [11] 972576-972662
BK063802 [11] 916414-916500
BK068299 [11] 986538-986624
IMGT000177 [9,10] 1083285-1083371
IMGT000207 753025-753111
M35650 [20] 69-99 *
M37277 [5] (3) 7949-7979 *
X97051 [1,5,13,15,18] 45954-46040
IGHD2IGHD2-8 IGHD2-8*02 F 14q32.33
RTPr
++
D1 J00233 [18] MAP 1-87 BK063800 [11] 983155-983241
BK068298 [11] 1005413-1005499
JADDFX010000192 [19] complement(126801-126887)
IGHD2IGHD2-15 IGHD2-15*01 F 14q32.33
RTPr
++
D2 J00234 [18] MAP 1-87 (4) AB019441 [12] 125668-125753
BK063799 [11] 981830-981916
BK063800 [11] 992409-992495
BK063801 [11] 937993-938079
BK063802 [11] 925668-925754
BK068298 [11] 1014665-1014751
BK068299 [11] 995792-995878
IMGT000207 762279-762365
JADDFX010000192 [19] complement(117670-117756)
X97051 [1,5,13,15,18] 55207-55293
IGHD2IGHD2-21 IGHD2-21*01 F 14q32.33
RTPr
++
D3 J00235 [18] MAP 1-84 BK063800 [11] 1001791-1001874
BK068298 [11] 1024047-1024130
JADDFX010000192 [19] complement(108291-108374)
IGHD2IGHD2-21 IGHD2-21*02 F 14q32.33
RTPr
++
D2-21 X97051 [1,5,13,15,18] MAP 64616-64699 AB019441 [12] 135075-135158
BK063799 [11] 991239-991322
BK063801 [11] 947402-947485
BK063802 [11] 935077-935160
BK068299 [11] 1005168-1005251
IMGT000207 771688-771771
IGHD3IGHD3-3 IGHD3-3*01 F 14q32.33
RTPr
++
DXP4 X13972 [5] MAP (3) 776-862 (4) AB019441 [12] 109265-109351
BK063799 [11] 965428-965514
BK063800 [11] 976008-976094
BK063802 [11] 909265-909351
BK068298 [11] 998266-998352
BK068299 [11] 979391-981579
IMGT000177 [9,10] 1076137-1076223
IMGT000207 745877-745963
JADDFX010000192 [19] complement(133948-134034)
M37277 [5] (3) 804-834 *
X97051 [1,5,13,15,18] 38806-38892
IGHD3IGHD3-3 IGHD3-3*02 F 14q32.33
RTPr
++
D23/7 X93618 [2] 2-88
IGHD3IGHD3-9 IGHD3-9*01 F 14q32.33
RTPr
++
DXP1 X13972 [5] MAP (3) 10447-10533 (4) AB019441 [12] 118944-119030
BK063799 [11] 975106-975192
BK063800 [11] 985685-985771
BK063801 [11] 931269-931355
BK063802 [11] 918944-919030
BK068298 [11] 1007941-1008027
BK068299 [11] 989068-989154
IMGT000207 755555-755641
JADDFX010000192 [19] complement(124271-124357)
M37277 [5] (3) 10475-10505 *
M37485 [4] 983-1013 *
D21/0,5 X93613 [2] 30-60 *
X97051 [1,5,13,15,18] 48484-48570
IGHD3IGHD3-10 IGHD3-10*01 F 14q32.33
RTPr
++
DXP'1 X13972 [5] MAP (3) 10631-10717 (4) AB019441 [12] 119128-119214
BK063799 [11] 975290-975376
BK063801 [11] 931453-931539
BK063802 [11] 919128-919214
BK068299 [11] 989252-991186
IMGT000207 755739-755825
M37277 [5] (3) 10659-10689 *
M37485 [4] 1166-1196 *
X97051 [1,5,13,15,18] 48668-48754
IGHD3IGHD3-10 IGHD3-10*02 F 14q32.33
RTPr
++
D21/7 X93615 [2] 2-87
IGHD3IGHD3-10 IGHD3-10*03 F 14q32.33
RTPr
++
BK063800 [11] MAP 985869-985955 BK068298 [11] 1008125-1008211
IGHD3IGHD3-16 IGHD3-16*01 F 14q32.33
RTPr
++
D21/10 X93614 [2] 2-94
IGHD3IGHD3-16 IGHD3-16*02 F 14q32.33
RTPr
++
X97051 [1,5,13,15,18] MAP 57524-57616 AB019441 [12] 127984-128076
BK063799 [11] 984147-984239
BK063801 [11] 940310-940402
BK063802 [11] 927985-928077
BK068299 [11] 998109-998201
IMGT000207 764596-764688
IGHD3IGHD3-16 IGHD3-16*03 F 14q32.33
RTPr
++
BK063800 [11] MAP 994726-994818 BK068298 [11] 1016982-1017074
JADDFX010000192 [19] complement(115347-115439)
IGHD3IGHD3-22 IGHD3-22*01 F 14q32.33
RTPr
++
D21/9 X93616 [2] MAP 2-88 (4) AB019441 [12] 137592-137678
BK063799 [11] 993756-993842
BK063800 [11] 1004260-1004346
BK063801 [11] 949919-950005
BK063802 [11] 937594-937680
BK068298 [11] 1026515-1026601
BK068299 [11] 1007637-1007723
JADDFX010000192 [19] complement(105977-106063)
X97051 [1,5,13,15,18] 67133-67219
IGHD4IGHD4-4 IGHD4-4*01 F 14q32.33
RTPr
++
DA4 X13972 [5] MAP (3) 1924-1995 (4) AB019441 [12] 110417-110488
BK063799 [11] 966580-966651
BK063800 [11] 977160-977231
BK063802 [11] 910417-910488
BK068298 [11] 999418-999489
BK068299 [11] 981504-981579
IMGT000177 [9,10] 1077289-1077360
IMGT000207 747029-747100
JADDFX010000192 [19] complement(132811-132882)
M37277 [5] (3) 1952-1967 *
X97051 [1,5,13,15,18] 39958-40029
IGHD4IGHD4-11 IGHD4-11*01 ORF (7) 14q32.33
RTPr
++
DA1 X13972 [5] MAP (3) 11522-11593 (4) AB019441 [12] 120024-120095
BK063799 [11] 976186-976257
BK063800 [11] 986765-986836
BK063801 [11] 932349-932420
BK063802 [11] 920024-920095
BK068298 [11] 1009021-1009092
BK068299 [11] F 991108-991186
IMGT000207 756635-756706
JADDFX010000192 [19] complement(123329-123400)
M37277 [5] (3) 11550-11565 *
X97051 [1,5,13,15,18] 49563-49634
IGHD4IGHD4-17 IGHD4-17*01 F 14q32.33
RTPr
++
D4-17 X97051 [1,5,13,15,18] MAP 58671-58742 AB019441 [12] 129131-129202
BK063799 [11] 985294-985365
BK063800 [11] 995873-995944
BK063801 [11] 941457-941528
BK063802 [11] 929132-929203
BK068298 [11] 1018129-1018200
BK068299 [11] 999256-999327
IMGT000207 765743-765814
JADDFX010000192 [19] complement(114221-114292)
IGHD4IGHD4-23 IGHD4-23*01 ORF (7) 14q32.33
RTPr
++
D4-23 X97051 [1,5,13,15,18] MAP 68306-68380 AB019441 [12] 138765-138839
BK063799 [11] 994929-995003
BK063800 [11] 1005433-1005507
BK063801 [11] 951092-951166
BK063802 [11] 938767-938841
BK068298 [11] 1027688-1027762
BK068299 [11] 1008810-1008884
JADDFX010000192 [19] complement(104816-104890)
IGHD5IGHD5-5 IGHD5-5*01 F 14q32.33
RTPr
++
DK4 X13972 [5] MAP (3) 2885-2960 (4) AB019441 [12] 111378-111453
BK063799 [11] 967541-967616
BK063800 [11] 978121-978196
BK063802 [11] 911378-911453
BK068298 [11] 1000379-1000454
BK068299 [11] 981504-981579
IMGT000177 [9,10] 1078250-1078325
IMGT000207 747990-748065
JADDFX010000192 [19] complement(131846-131921)
M37277 [5] (3) 2913-2932 *
X97051 [1,5,13,15,18] 40919-40994
IGHD5IGHD5-12 IGHD5-12*01 F 14q32.33
RTPr
++
DK1 X13972 [5] MAP (3) 12478-12556 (4) AB019441 [12] 120984-121062
BK063799 [11] 977146-977224
BK063800 [11] 987725-987803
BK063801 [11] 933309-933387
BK063802 [11] 920984-921062
BK068298 [11] 1009981-1010059
BK068299 [11] 991108-991186
IMGT000207 757595-757673
JADDFX010000192 [19] complement(122362-122440)
M37277 [5] (3) 12506-12528 *
X97051 [1,5,13,15,18] 50523-50601
IGHD5IGHD5-18 IGHD5-18*01 F 14q32.33
RTPr
++
D5-18 X97051 [1,5,13,15,18] MAP 59633-59708 AB019441 [12] 130093-130168
BK063799 [11] 986256-986331
BK063801 [11] Unknown
BK063802 [11] 930094-930169
BK068299 [11] 1000218-1000293
IMGT000207 766705-766780
IGHD5IGHD5-18 IGHD5-18*02 ORF (1) 14q32.33
RTPr
++
BK063800 [11] MAP 996834-996912 BK068298 [11] 1019090-1019168
JADDFX010000192 [19] complement(113253-113331)
IGHD5IGHD5-24 IGHD5-24*01 ORF (8) 14q32.33
RTPr
++
D5-24 X97051 [1,5,13,15,18] MAP 69272-69347 AB019441 [12] 139730-139805
BK063799 [11] 995895-995970
BK063800 [11] 1006399-1006474
BK063801 [11] 952058-952133
BK063802 [11] 939733-939808
BK068298 [11] 1028654-1028729
BK068299 [11] 1009776-1009851
JADDFX010000192 [19] complement(103849-103924)
IGHD6IGHD6-6 IGHD6-6*01 F 14q32.33
RTPr
++
DN4 X13972 [5] MAP (3) 4734-4807 (4) AB019441 [12] 113227-113300
BK063799 [11] 969390-969463
BK063800 [11] 979970-980042
BK063802 [11] 913228-913300
BK068298 [11] 1002228-1002300
BK068299 [11] 983353-983425
IMGT000177 [9,10] 1080098-1080171
IMGT000207 ORF (9) 749839-749912
JADDFX010000192 [19] complement(130000-130072)
M37277 [5] (3) 4762-4779 *
X97051 [1,5,13,15,18] 42767-42840
IGHD6IGHD6-13 IGHD6-13*01 F 14q32.33
RTPr
++
DN1 X13972 [5] MAP (3) 13983-14059 (4) AB019441 [12] 122493-122569
BK063799 [11] 978655-978731
BK063800 [11] 989234-989310
BK063801 [11] 934818-934894
BK063802 [11] 922493-922569
BK068298 [11] 1011490-1011566
BK068299 [11] 992617-992693
IMGT000207 759104-759180
JADDFX010000192 [19] complement(120855-120931)
M37277 [5] (3) 14011-14031 *
X97051 [1,5,13,15,18] 52032-52108
IGHD6IGHD6-19 IGHD6-19*01 F 14q32.33
RTPr
++
D6-19 X97051 [1,5,13,15,18] MAP 61475-61551 AB019441 [12] 131934-132010
BK063799 [11] 988098-988174
BK063800 [11] 998683-998759
BK063801 [11] 944261-944337
BK063802 [11] 931936-932012
BK068298 [11] 1020939-1021015
BK068299 [11] 1002060-1002136
IMGT000207 768547-768623
JADDFX010000192 [19] complement(111406-111482)
IGHD6IGHD6-25 IGHD6-25*01 F 14q32.33
RTPr
++
D6-25 X97051 [1,5,13,15,18] MAP 71638-71711 AB019441 [12] 142095-142168
BK063799 [11] 998261-998334
BK063800 [11] 1008765-1008810
BK063801 [11] 954424-954497
BK063802 [11] 942099-942172
BK068298 [11] 1031020-1031093
BK068299 [11] 1012142-1012215
JADDFX010000192 [19] complement(101485-101558)
IGHD7IGHD7-27 IGHD7-27*01 F 14q32.33
RTPr
++
DHQ52 J00256 [3,14,16] MAP 592-659 (4) BK063799 [11] 1014256-1014322
BK063800 [11] 1024348-1024414
BK063801 [11] 970024-970091
BK063802 [11] 957742-957808
BK068298 [11] 1046603-1046662
BK068299 [11] 1028128-1028195
JADDFX010000192 [19] complement(86486-86553)
X86355 [13] 154-220
X86356 [13] 154-220
X86357 [13] 154-220
X86358 [13] Unknown
X86359 [13] 154-220
X97051 [1,5,13,15,18] 87431-87497

✤ : NCBI accession number that correspond to a previously internal IMGT accession number. See the correspondence table.

IMGT notes:
  1. noncanonical 5'D-NONAMER: ggttatcgt instead of ggattttgt
  2. The IMGT000054 sequence is in direct orientation relative to the IMGT Homo sapiens IGH locus. They is reverse-complement of the GEDI (GenBank/ENA/DDBJ/IMGT/LIGM-DB) sequence AC279151, which is in opposite orientation relative to the IMGT Homo sapiens IGH locus.
  3. The original sequence is X13972 (14928 bp), M37277 (14929 bp) secondary accession number presents an INSERTION of one nucleotide C between positions 14587/14588 (in X13972).
  4. Arbitrary limits of the DNA sequence assigned to the different D-GENEs. D-GENE positions are those in the Accession number X97051.
  5. ORF, because unusual 5'D HEPTAMER CACAGCG instead of CACAGTG.
  6. INSERTION of one nucleotide C between positions 14587/14588 is after the coding region of the IGHD1-14, and do not concern the recombination signals either.
  7. ORF, because unusual 5'D HEPTAMER TGCTGTG instead of CACAGTG.
  8. ORF, because unusual 5'D HEPTAMER GGCCGTG instead of CACAGTG.
  9. noncanonical 5'D-NONAMER: agtttctga instead of ggtttttgt
IMGT references:
  1. Buluwela L. et al., A VH gene is located within 95 Kb of the human immunoglobulin heavy chain constant region genes, Eur. J. Immunol, vol. 18, no. 11, 1988, pp. 1843-1845. PUBMED: 3144456
  2. Buluwela L. et al., The use of chromosomal translocations to study human immunoglobulin gene organization: mapping DH segments within 35 kb of the C mu gene and identification of a new DH locus, EMBO J, vol. 7, no. 7, 1988, pp. 2003-2010. PUBMED: 3138112
  3. Flanagan J.G. et al., The sequence of a human immunoglobulin epsilon heavy chain constant region gene, and evidence for three non-allelic genes, EMBO J, vol. 1, no. 5, 1982, pp. 655-660. PUBMED: 6234164
  4. Huang C. et al., Analysis of members of the Dxp family of human immunoglobulin diversity gene segments, Unpublished.
  5. Ichihara Y. et al., Organization of human immunoglobulin heavy chain diversity gene loci, EMBO J, vol. 7, no. 13, 1988, pp. 4141-4150. PUBMED: 3243276
  6. Li,H. et al., Direct Submission, Genomics Institute, UCSC Genomics Institute, 1156 High Street, Santa Cruz, CA 95064, USA.
  7. Li,H. et al., Human Pangenomics Reference Consortium: Year 1 Assemblies, Unpublished.
  8. Liao,W.W. et al., A draft human pangenome reference, Nature, vol. 617, no. 7960, 2023, pp. 312-324.
  9. Liao W.W. et al., A draft human pangenome reference, Nature, vol. 617, no. 7960, 2023, pp. 312-324. PUBMED: 37165242
  10. Li H. et al., Human Pangenomics Reference Consortium: Year 1 Assemblies, Unpublished.
  11. Manso,T. et al., IMGT(R) databases, related tools and web resources through three main axes of research and development, Nucleic Acids Res, vol. 50, no. D1, 2022, pp. D1262-D1272. PUBMED: 34875068
  12. Matsuda F. et al., The complete nucleotide sequence of the human immunoglobulin heavy chain variable region locus, J. Exp. Med, vol. 188, no. 11, 1998, pp. 2151-2162. DOI: 10.1084/jem.188.11.2151
  13. Mattila P.S. et al., Extensive allelic sequence variation in the J region of the human immunoglobulin heavy chain gene locus, Eur. J. Immunol, vol. 25, no. 9, 1995, pp. 2578-2582. PUBMED: 7589129
  14. Rabbitts T.H. et al., Human immunoglobulin heavy chain genes: evolutionary comparisons of C mu, C delta and C gamma genes and associated switch sequences, Nucleic Acids Res, vol. 9, no. 18, 1981, pp. 4509-4524. DOI: 10.1093/nar/9.18.4509
  15. Rabbitts T.H. et al., Transcription enhancer identified near the human C mu immunoglobulin heavy chain gene is unavailable to the translocated c-myc gene in a Burkitt lymphoma, Nature, vol. 306, no. 5945, 1983, pp. 806-809. DOI: 10.1038/306806a0
  16. Ravetch J.V. et al., Structure of the human immunoglobulin mu locus: characterization of embryonic and rearranged J and D genes, Cell, vol. 27, no. 3 Pt 2, 1981, pp. 583-591. DOI: 10.1016/0092-8674(81)90400-1
  17. Rodriguez O.L. et al., An ethnically diverse genomic reference resource for human immune receptor gene loci, Unpublished.
  18. Siebenlist U. et al., Human immunoglobulin D segments encoded in tandem multigenic families, Nature, vol. 294, no. 5842, 1981, pp. 631-635. DOI: 10.1038/294631a0
  19. Wong,K. et al., Towards a reference genome that captures global genetic diversity, Unpublished.
  20. Zong S.Q. et al., Human immunoglobulin D segments: isolation of a new D segment and polymorphic deletion of the D1 segment, Immunol. Lett, vol. 17, no. 4, 1988, pp. 329-333. DOI: 10.1016/0165-2478(88)90006-5