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ß-Lactamase Classification and Amino Acid Sequences for TEM, SHV and OXA Extended-Spectrum and Inhibitor Resistant Enzymes

NOTICE 6/2/2015: This website will be transitioning at the end of June to a new location with new requirements for numbered allele assignment. As of July 1, the Lahey site is no longer assigning beta-lactamase allele numbers. Please contact https://www.ncbi.nlm.nih.gov/pathogens/submit-beta-lactamase for further details
These tables grew out of an attempt to standardize the nomenclature for the growing number of TEM ß-lactamases and the intent to make it easier for investigators to discover if particular amino acid substitutions had been described before (1). To the TEM table we have added tables for SHV and OXA enzymes with extended-spectrum or other properties and have provided references to sources for sequence information. As of May, 2001, we have added a listing for other rapidly growing enzyme groups, including OXA-, CTX-M-, CMY-, AmpC, CARB-, IMP-, VIM-, KPC-, GES-, PER-, VEB-, and other types. Additional data about AmpC-type ß-lactamases can be found in (2). Another web site for TEM, SHV, and class B enzymes can be found at http://www.laced.uni-stuttgart.de. A web site for LEN, OXY, and OKP ß-lactamases is located at http://bigsdb.web.pasteur.fr/klebsiella/klebsiella.html. More references about the ß-lactamases themselves can be found in reviews by Bush, Jacoby, & Medeiros (3), Livermore (4), Bradford (5), Jacoby & Munoz-Price (6), Paterson & Bonomo (7), Queenan & Bush (8), Poirel, Naas, & Nordmann (13), and Bush & Jacoby (12). Structural considerations have been reviewed by Knox (9). Residue numbering for TEM and SHV enzymes follows Ambler et al. (10). More on ß-lactamase nomenclature can be found in (11).
To qualify for a new number the enzyme must have been found in nature and not created in the laboratory. The amino acid sequence of the native enzyme and not merely the nucleotide sequence should be unique. Changes in the promoter sequence do not qualify as new derivatives. Substitutions in the signal sequence do not alter the mature protein but since they may provide a useful epidemiological marker and have already been used to define new varieties, such changes are considered valid. We will accept any amino acid alteration as defining a new variety, even if no functional change is currently evident, since most enzymes will have been sequenced because of a change in properties and an effect on interaction with a future substrate or inhibitor is always possible. We will also assign numbers sequentially independent of ß-lactamase properties. Application for a new enzyme number can be made to either of us. With the investigator's permission the new sequence can be made public before publication so that the information is available without delay.
The TEM and SHV tables can be downloaded as Excel files by clicking on the Excel logo next to the table title.

Karen Bush Indiana University, Bloomington, IN 47405 Email: karbush@indiana.edu
Timothy Palzkill Baylor College of Medicine, Houston, TX 77030 Email: timothyp@bcm.edu
George Jacoby Lahey Clinic, Burlington, MA 01805 Email: gajacoby50@gmail.com

1. Bush, K. and G. Jacoby. Nomenclature of TEM ß-lactamases. 1997. J. Antimicrob. Chemother. 39:1-3.
2. Jacoby, G.A. AmpC ß-Lactamases. 2009. Clin. Microbiol. Rev. 22:161-182.
3. Bush, K., G.A. Jacoby, and A.A. Medeiros. 1995. A functional classification scheme for ß-lactamases and its correlation with molecular structure. Antimicrob. Agents Chemother. 39:1211-1233.
4. Livermore, D. M. 1995. ß-Lactamases in laboratory and clinical resistance. Clin. Microbiol. Rev. 8:557-584.
5. Bradford, P. A. 2001. Extended-spectrum ß-lactamases in the 21st century: characterization, epidemiology, and detection of this important resistance threat. Clin. Microbiol. Rev. 14:933-951.
6. Jacoby, G.A., and Munoz-Price, L.S. 2005. The new ß-lactamases. New Engl. J. Med. 352:380-391.
7. Paterson, D. L., and R. A. Bonomo. 2005. Extended-spectrum ß-lactamases: a clinical update. Clin. Microbiol. Rev. 18:657-686.
8. Queenan, A.M., and K. Bush. 2007. Carbapenemases: the versatile ß-lactamases. Clin. Microbiol. Rev. 20:440-458.
9. Knox, J.R. 1995. Extended-spectrum and inhibitor-resistant TEM-type ß-lactamases: mutation, specificity, and three-dimensional structure. Antimicrob. Agents Chemother. 39:2593-2601.
10. Ambler, R.P., A.F.W. Coulson, J.-M. Frère, J.-M. Ghuysen, B. Joris, M. Forsman, R.C. Levesque, G. Tiraby, and S.G. Waley. 1991. A standard numbering scheme for the class A ß-lactamases. Biochem. J. 276:269-272.
11. Jacoby, G.A. 2006. ß-Lactamase nomenclature. Antimicrob Agents Chemother 50:1123-1129.
12. Bush, K, and G.A. Jacoby. 2010. Updated functional classification of ß-lactamases. Antimicrob. Agents. Chemother. 54:969-976.
13. Poirel, L., T. Naas, and P. Nordmann. 2010. Diversity, epidemiology, and genetics of class D ß-lactamases. Antimicrob. Agents Chemother. 54:24-38.
14. Evans, B.A., and S.G.B.Amyes. 2014. OXA Beta-lactamases. Clin. Microbiol. Rev. 27:241-253.

SHV Extended-Spectrum and Inhibitor Resistant ß-Lactamases

Amino acid at position
ß-Lactamase GenBank Accession Number Alternate Name Phenotype 5 7 8 10 11 12 13 14 18 19 20 22 24 25 31 35 38 39 40 43 47 51 52 53 54 60 61 64 65 69 70 71 75 79 80 87 89 91 98 101 104 106 112 113 114 119 122 123 124 126 129 130 134 136 140 141 142 144 145 146 147 148 149 150 151 154 156 157 158 164 169 172 173 174 175 179 184 187 188 192 193 194 195 196 202 203 205 206 211 213 214 215 226 234 235 238 240 241 242 243 248 250 252 254 255 256 258 260 262 268 269 271 273 274 275 276 278 282 286 287 292 pI Reference
SHV-1 AF148850 2b M Y I L C I I S T L P A H A E L S Q L R I L A S G W R E R M S T V A V G E L R D D S H L A V L C A A M S A N A T V G P A G L T A F Q G D N R L A L P G D A A A K L L T S R S R Q M D D R P K T G E R G A A L P N N K E I V T P S A E R N Q I L I R 7.6 1
SHV-2 AF148851 2be S 7.6 1
SHV-2A X98102 2be Q S 7.6 2
SHV-3 P30896 (protein) 2be L S 7 3
SHV-4 2be L S K 7.8 4
SHV-5 X55640 2be S K 8.2
SHV-6 2be A 7.6 6
SHV-7 U20270 2be F S S K 7.6 7
SHV-8 U92041 2be N 7.6 8
SHV-9 SHV-5a 2be Del R N V S K 8.2 9
SHV-10 2br Del G R N V S K 8.2 10
SHV-11 X98101 SHV-1-2a 2b Q 7.6 11
SHV-12 AJ920369 SHV-5-2a 2be Q S K 8.2 11
SHV-13 AF164577 2be Q A 7.6 12
SHV-14 AF226622 2b F S 7 13
SHV-15 2be Q M K S K 14
SHV-16 163DRWET167 insertion 2be 7.6 15
SHV-17 Withdrawn
SHV-18 AF132290 2be F S A K 7.8 16
SHV-19 2b F 7.6 17
SHV-20 2b F S 7.6 17
SHV-21 2b F F S 17
SHV-22 2b K S K 7.6 17
SHV-23 ?2be F G S K 21
SHV-24 AB023477 2be G 18
SHV-25 2b A Q V 7.5 19
SHV-26 KF585144 2br T 7.6 19
SHV-27 2be D 22
SHV-28 ? F 23
SHV-29 ? Q S A 24
SHV-30 AY661885 2be F S S 25
SHV-31 AY277255 2be Q K 26
SHV-32 AY037778 2b V D 27
SHV-33 2b S 27
SHV-34 AY036620 2be F S G S 28
SHV-35 AY070258 ? Q K G 29
SHV-36 AF467947 ? V Q
SHV-37 AF467948 ? A Q V H 41
SHV-38 AY079099 2be V 7.6 30
SHV-39 AY150585 Invalidated as only a partial sequence. 2be S S 31
SHV-40 AF535128 2be Q G 34
SHV-41 AF535129 2be F 34
SHV-42 AF535130 2be S V 34
SHV-43 AY065991 2b F S 8 37
SHV-44 AY259119 2b L 38
SHV-45 AF547625 2be D S K 8.2 39
SHV-46 AY210887 2be N S K 40
SHV-47 Withdrawn
SHV-48 AY263404 2b I 7.6 42
SHV-49 AY528718 2br I 7.6 43
SHV-50 AY288915 ? T H 44
SHV-51 AY289548 ? L A 45
SHV-52 HQ845196 ? A Q I I 46
SHV-53 AY590467 ? Q I 47
SHV-54 Not yet released F 48
SHV-55 DQ054528 2be F S K 49
SHV-56 EU586041 2br Q R 50
SHV-57 AY223863 2be R 51
SHV-58 Not yet released
SHV-59 AY790341 ? V L 52
SHV-60 AJ866283 2b Q T 7.6 57
SHV-61 AJ866284 2b R Q 7.6 57
SHV-62 AJ866285 2b Q Y 7.4 57
SHV-63 EU342351 ? E N 53
SHV-64 DQ174304 2be Q L S K 60
SHV-65 DQ174305 ? T 61
SHV-66 DQ174306 2be Q Q S K 62
SHV-67 DQ174307 ? Q C 63
SHV-68 Not yet released 55
SHV-69 DQ174308 ? Q T 64
SHV-70 DQ013287 2be Q V 56
SHV-71 AM176546 2b Y V 7.6 57
SHV-72 AM176547 2br F V R 7.6 65a
SHV-73 AM176548 2b Del V R 7.6 57
SHV-74 AM176549 2b T Q 7.6 57
SHV-75 AM176550 2b H 7.6 57
SHV-76 AM176551 2b A Q E 7.6 57
SHV-77 AM176552 2b Q A 7.6 57
SHV-78 AM176553 2b S 7.6 57
SHV-79 AM176554 2b Q V 7.6 57
SHV-80 AM176555 2b Q T 7.6 57
SHV-81 AM176556 2b Q S 7.6 57
SHV-82 AM176557 2b V T Q 7.6 57
SHV-83 AM176558 2b K 7.6 57
SHV-84 2br R 65
SHV-85 DQ322460 2b M Q 66
SHV-86 DQ328802 2be Q S R 67
SHV-87 Not yet released 68
SHV-88 Not yet released 68
SHV-89 DQ193536 2b Q V 69
SHV-90 2be T S K 8.2 70
SHV-91 2be S K 7.6 70
SHV-92 EF836922 ? Q I I 7.6 71
SHV-93 EF373969 ? Q T D 72
SHV-94 EF373970 ? Q H 74
SHV-95 EF373972 ? L 75
SHV-96 EF373971 ? Q A 76
SHV-97 EF373973 ? K S 77
SHV-98 AM941844 2be I 73
SHV-99 AM941845 2be G 73
SHV-100 AM941846 35SESQLSGRVGMIE36 insertion 2be 73
SHV-101 EU155018 ? G 78
SHV-102 EU024485 2be A 79
SHV-103 EU032604 ? R 80
SHV-104 EU274581 2be S 81
SHV-105 FJ194944 2be F S D S K 82
SHV-106 AM922307 ?2be F S 7.6 57
SHV-107 AM922308 2br Q A 7.6 57
SHV-108 AM922309 ? Q I 7.6 57
SHV-109 EU418913 2b R Q M 84
SHV-110 HQ877615 ? Q D 85
SHV-111 ? S 85
SHV-112 ? D 87
SHV-113 ? K 88
SHV-114 ? V Q V 88
SHV-115 2be H K K 88
SHV-116 ? H 89
SHV-117 ? L 89
SHV-118 Not yet released 90
SHV-119 KJ776406 ? H Q Q 91
SHV-120 JF812965 ?