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Main TEM SHV OXA ESBL Others QNR

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 ? K 91
SHV-121 GQ428198 ? A V Q V 92
SHV-122 GQ290211 ? Q A S 93
SHV-123 GQ390805 ? Q S K G I 94
SHV-124 GQ390806 ? D S K 94
SHV-125 GQ390807 ? N 94
SHV-126 GQ390808 ? K T 94
SHV-127 GQ390809 ? F N M 94
SHV-128 GU932590 2be Q R S K T P 106
SHV-129 GU827715 2be Q S K L D 95
SHV-130 withdrawn
SHV-131 withdrawn
SHV-132 Not yet released
SHV-133 AB551737 ? G 98
SHV-134 HM559945 2be Q E S K 99
SHV-135 HQ637576 ? K Q T 100
SHV-136 HQ661362 Identical to SHV-121 A V Q V 101
SHV-137 HQ661363 A H 102
SHV-138 Not yet released 103
SHV-139 Not yet released 103
SHV-140 JN051143 2b K S 104
SHV-141 JQ388884 2b S S 105
SHV-142 JQ029959 A Q 106
SHV-143 JQ341060 L 107
SHV-144 JQ926986 Q V 108
SHV-145 JX013655 R 109
SHV-146 KP868752 K K 110
SHV-147 JX121114 A S 111
SHV-148 JX121115 Q 111
SHV-149 JX121116 R S S 111
SHV-150 JX121117 V 111
SHV-151 JX121118 P S 111
SHV-152 JX121119 R S 111
SHV-153 JX121120 S V 111
SHV-154 JX121121 S S K 111
SHV-155 JX121122 Q Q 111
SHV-156 JX121123 Q P T 111
SHV-157 JX121124 Q M 111
SHV-158 JX121125 Q A 111
SHV-159 JX121126 Q G 111
SHV-160 JX121127 Q G S K 111
SHV-161 JX121128 S 111
SHV-162 JX121129 S V S 111
SHV-163 JX121130 S S Q 111
SHV-164 HE981194 2b Q 112
SHV-165 JX121131 P S K 111
SHV-166 Not yet released 113
SHV-167 AB733453 D Y V 114
SHV-168 JX870080 F V 115
SHV-169 2b H 116
SHV-170 2b F L 116
SHV-171 2b F F 116
SHV-172 KF513177 Q D 117
SHV-173 KF513178 Q Q 118
SHV-174 Not yet released Q V A 119
SHV-175 Y 120
SHV-176 Y W 120
SHV-177 Y W D 120
SHV-178 KF705209 Q V 121
SHV-179 KF705208 N 122
SHV-180 KP050487 H T 122
SHV-181 KP050488 Q S 122
SHV-182 KP050489 Q T 122
SHV-183 HG934764 Insertion A between 186 & 187 2be Q S K 123
SHV-184 Q P 124
SHV-185 KM233164 M 125
SHV-186 KM233165 T C 125
SHV-187 LN515533 Start VVKRY... 126
SHV-188 LN515534 Insertion 169KL170, start VVKRY... 126
SHV-189 KP050494 F S R 127
SHV-190 KP868753 Q L 128
SHV-191 KP868754 V D 128
SHV-192 Not yet released F 129
SHV-193 KR347170 Q L 128
OXA Extended-Spectrum ß-Lactamases (Incomplete)

B-Lactamase Alternate Name Amino acid at position
5 20 58 76 110 127 146 149 164 165 167 169 184 241 258 272 pI References
OXA-10 (parent) PSE-2 I G D N T A N W L G Y E S E 6.1 1
OXA-11 S D 6.4 2
OXA-14 D 6.2 3
OXA-16 T D 6.2 5
OXA-17 S 6.1 9
OXA-19 T S N S D F G N A 7.5 8
OXA-28 T S N S G F G N A 8.1 11
OXA-35 T S N S F G N A 8 14
OXA-142 S D 6.3 16
OXA-145 T S N S del W F G N A 8.7 16
OXA-147 T S N S L F G N A 8.0 17
OXA-2 (parent) N D W G L 7.7 6
OXA-15 G 8 7
OXA-32 I 9.0 15
OXA-34 C 8.9 16
OXA-36 Y 9.2 16
OXA-141 S 9.1 16
OXA-161 D 18
OXA-226 R 19
The OXA enzyme numbering follows the DBL system of Couture, F., J. Lachapelle, and R. C. Levesque. 1992. Phylogeny of LCR-1 and OXA-5 with class A and class D ß-lactamases. Mol. Microbiol. 6:1693-1705.
SHV References

1. Bradford, P.A. 1999. Automated thermal cycling is superior to traditional methods for nucleotide sequencing of blaSHV genes. Antimicrob. Agents Chemother. 43: 2960-2963.
2. Podbielski, A., J. Schonling, B. Melzer, K. Warnatz, and H.-G. Leusch. 1991. Molecular characterization of a new plasmid-encoded SHV-type ß-lactamase (SHV-2 variant) conferring high-level cefotaxime resistance upon Klebsiella pneumoniae. J. Gen. Microbiol. 137:569-578.
3. Nicolas, M.-H., V. Jarlier, N. Honore, A. Philippon, and S. T. Cole. 1989. Molecular characterization of the gene encoding SHV-3 ß-lactamase responsible for transferable cefotaxime resistance in clinical isolates of Klebsiella pneumoniae. Antimicrob. Agents Chemother. 33:2096-2100.
4. Peduzzi, J., M. Barthelemy, K. Tiwari, D. Mattioni, and R. Labia. 1989. Structural features related to hydrolytic activity against ceftazidime of plasmid-mediated SHV-type CAZ-5 ß-lactamase. Antimicrob. Agents Chemother. 33:2160-2163.
5. Billot-Klein, D., L. Gutmann, and E. Collatz. 1990. Nucleotide sequence of the SHV-5 ß-lactamase gene of a Klebsiella pneumoniae plasmid. Antimicrob. Agents Chemother. 34:2439-2441.
6. Arlet, G., M. Rouveau, and A. Philippon. 1997. Substitution of alanine for aspartate at position 179 in the SHV-6 extended-spectrum ß-lactamase. FEMS Microbiol. Lett. 152:163-167.
7. Bradford, P. A., C. Urban, A. Jaiswal, N. Mariano, B. A. Rasmussen, S. J. Projan, J. J. Rahal, and K. Bush. 1995. SHV-7, a novel cefotaxime-hydrolyzing ß-lactamase, identified in Escherichia coli isolated from hospitalized nursing home patients. Antimicrob. Agents Chemother. 39:899-905.
8. Rasheed, J. K., C. Jay, B. Metchock, F. Berkowitz, L. Weigel, J. Crellin, C. Steward, B. Hill, A. A. Medeiros, F. C. Tenover. 1997. Evolution of extended-spectrum ß-lactam resistance (SHV-8) in a strain of Escherichia coli during multiple episodes of bacteremia. Antimicrob. Agents Chemother. 41:647-653.
9. Prinarakis, E. E., E. Tzelepi, M. Gazouli, A. F. Mentis, and L. S. Tzouvelekis. 1996. Characterization of a novel SHV ß-lactamase variant that resembles the SHV-5 enzyme. FEMS Microbiol. Lett. 139:229-234.
10. Prinarakis, E. E., V. Miriagou, E. Tzelepi, M. Gazouli, and L. S. Tzouvelekis. 1997. Emergence of an inhibitor-resistant ß-lactamase (SHV-10) derived from an SHV-5 variant. Antimicrob. Agents Chemother. 41:838-840.
11. Nuesch-Inderbinen, M.T., F.H. Kayser, and H. Hachler. 1997. Survey and molecular genetics of SHV ß-lactamases in Enterobacteriaceae in Switzerland: two novel enzymes, SHV-11 and SHV-12. Antimicrob. Agents Chemother. 41:943-949.
12. Yuan, M., L.M.C. Hall, P.H.M. Savelkoul, C.M.J.E. Vandenbroucke-Grauls, and D. M. Livermore. 2000. SHV-13, a novel extended-spectrum ß-lactamase, in Klebsiella pneumoniae isolates from patients in an intensive care unit in Amsterdam. Antimicrob. Agents Chemother. 44:1081-1084.
13. Yuan, M., L.M.C. Hall, J. Hoogkamp-Korstanje, and D.M. Livermore. 2001. SHV-14, a novel ß-lactamase variant in Klebsiella pneumoniae form Nijmegen, The Netherlands. Antimicrob. Agents Chemother. 45: 309-311.
14. Child, J.A., D.M. Livermore, and M.J. Gill. SHV-15 and SMY-2 extended spectrum ß-lactamases in a strain of Escherichia coli imported from India into the United Kingdom. GenBank AJ011428
15. Arpin, C., R. Labia, C. Andre, C. Frigo, Z. El Harrif, and C. Quentin. 2001. SHV-16, a ß-lactamase with a pentapeptide duplication in the omega loop. Antimicrob. Agents Chemother. 45: 2480-2485.
16. Rasheed, J.K., G.J. Anderson, H. Yigit, A.M. Queenan, A. Doménech-Sanchez, J.M. Swenson, J.W. Biddle, M.J. Ferraro, G.A. Jacoby, and F.C. Tenover. 2000. Characterization of the extended-spectrum ß-lactamase reference strain, Klebsiella pneumoniae K6 (ATCC 700603), which produces the novel enzyme SHV-18. Antimicrob. Agents Chemother. 44:2382-2388.
17. Essack, S.Y., L.M.C. Hall, D.G. Pillary, M.L. McFadyen, and D.M. Livermore. 2001. Complexity and diversity of Klebsiella pneumoniae strains with extended-spectrum ß-lactamases isolated in 1994 and 1996 at a teaching hospital in Durban, South Africa. Antimicrob. Agents Chemother. 45: 88-95.
18. Kurokawa, H., T. Yagi, N. Shibata, K. Shibayama, K. Kamachi, and Y. Arakawa. 2000. A new SHV-derived extended-spectrum ß-lactamase (SHV-24) that hydrolyzed ceftazidime through a single-amino-acid substitution (D179G) in the omega-loop. Antimicrob. Agents Chemother. 44:1725-1727.
19. Chang, F.-Y., L.K Siu, C.-P. Fung, M.-H. Huang, and M. Ho. 2001. Diversity of SHV and TEM ß-lactamases in Klebsiella pneumoniae: gene evolution in Northern Taiwan and two novel ß-lactamases, SHV-25 and SHV-26. Antimicrob. Agents Chemother. 45: 2407-2413.
21. Essack, S. Y., L. M. Hall, and D. M. Livermore. 2004. Klebsiella pneumoniae isolate from South Africa with multiple TEM, SHV and AmpC ß-lactamases. Int. J. Antimicrob. Agents. 23:398-400.
22. Corkill, J.E., L.E. Cuevas, R.Q. Gurgel, J. Greensill, and C.A. Hart. 2001. SHV-27, a novel cefotaxime-hydrolyzing ß-lactamase, identified in Klebsiella pneumoniae isolates from a Brazilian hospital. J. Antimicrob. Chemother. 47: 463-465.
23. Yu,Y.,Zhou,W. and Chen,Y.GenBank AF299299
24. Yigit, H., A.M. Queenan, G.J. Anderson, A. Domenech-Sanchez, J.W. Biddle, C.D. Steward, S. Alberti, K. Bush, and F. Tenover. 2001. Novel carbapenem-hydrolyzing ß-lactamase KPC-1, from a carbapenem-resistant strain of Klebsiella pneumoniae. Antimicrob. Agents Chemother. 45: 1151-1161.
25. Szabó, D., M. A. Melan, A. M. Hujer, R. A. Bonomo, K. M. Hujer, C. R. Bethel, K. Kristõf, and D. L. Paterson. 2005. Molecular analysis of the simultaneous production of two SHV-type extended-spectrum beta-lactamases in a clinical isolate of Enterobacter cloacae by using single-nucleotide polymorphism genotyping. Antimicrob. Agents Chemother. 49:4716-4720.
26. Mazzariol, A., E. Roelofsen, R. Koncan, A. Voss, and G. Cornaglia. 2007. Detection of a new SHV-type extended-spectrum ß-lactamase, SHV-31, in a Klebsiella pneumoniae strain causing a large nosocomial outbreak in The Netherlands. Antimicrob. Agents Chemother. 51:1082-1084.
27. Chaves, J., M. G. Ladona, C. Segura, A. Coira, R. Reig, and C. Ampurdanés. 2001. SHV-1 ß-lactamase is mainly a chromosomally encoded species-specific enzyme in Klebsiella pneumoniae. Antimicrob. Agents Chemother. 45:2856-61.
28. Heritage, J., P.A. Chambers, C. Tyndall, and E.S. Buescher. 2003. SHV-34: an extended-spectrum ß-lactamase encoded by an epidemic plasmid. J. Antimicrob. Chemother. 52:1015-1017.
29. Alobwede,I.N., M'Zali,F.H., Todd,N., Hancox,A., Allen,M.,Heritage,J. and Hawkey,P.M.GenBank AY070258
30. Poirel, L., C. Héritier, I. Podglajen, W. Sougakoff, L. Gutmann, and P. Nordmann. 2003. Emergence in Klebsiella pneumoniae of a chromosome-encoded SHV ß-lactamase that compromises the efficacy of imipenem. Antimicrob. Agents Chemother. 47:755-758.
31. Bello,H.M., Dominguez,M.P., Dashti,A.A., Gonzalez-Rocha,G.E. and Amyes,S.G.B. GenBank AY150585. However, since the sequence in AY150585 is only 285 nucleotidew or 95 amino acids long, SHV-39 is not adequately described to remain a valid designation.
34. Mulvey, M. R., E. Bryce, D. Boyd, M. Ofner-Agostini, S. Christianson, A. E. Simor, and S. Paton. 2004. Ambler class A extended-spectrum beta-lactamase-producing Escherichia coli and Klebsiella spp. in Canadian hospitals. Antimicrob Agents Chemother 48:1204-14.
37. Wang, H., S. Kelkar, W. Wu, M. Chen, and J. P. Quinn. 2003. Clinical isolates of Enterobacteriaceae producing extended-spectrum ß-lactamases: prevalence of CTX-M-3 at a hospital in China. Antimicrob. Agents Chemother. 47:790-793.
38. Arpin, C., V. Dubois, L. Coulange, C. André, I. Fischer, P. Noury, F. Grobost, J-P. Brochet, J. Jullin, B. Dutilh, G. Larribet, I. Lagrange, and C. Quentin. 2003. Extended-spectrum ß-lactamase-producing Enterobacteriaceae in community and private health care centers. Antimicrob. Agents Chemother. 47:3506-3514, 2003.
39. Corkill,J.E., Irala,J.D. and Charles,H.A. GenBank AF547625.
40. Yigit, H., A.M. Queenan, J.K. Rasheed, J.W. Biddle, A. Domenech-Sanchez, S. Alberti, K. Bush, and F.C. Tenover. 2003. Carbapenem-resistant strain of Klebsiella oxytoca harboring carbapenem-hydrolyzing ß-lactamase KPC-2. Antimicrob. Agents Chemother. 47: 3881-3889, 2003.
41. Alobwede,I.N., M'Zali,F.H., Todd,N., Hancox,A., Allen,M.,Heritage,J. and Hawkey,P.M. GenBank AF467948.
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43. Dubois, V, L. Poirel, C. Arpin, L. Coulange, C. Bebear, P, Nordmann, and C. Quentin. 2004. SHV-49, a novel inhibitor-resistant ß-lactamase in a clinical isolate of Klebsiella pneumoniae. Antimicrob. Agents Chemother: 48:4466-4469, 2004.
44. Boyd, D.A. and M.R. Mulvey. GenBank AY288915.
45. Boyd, D.A. and M.R. Mulvey. GenBank AY289548.
46. Obert, C.A, Gordon, D.M., and Riley, M.A.. GenBank AY265888 (partial sequence).
47. Obert,C.A., Riley,M.A. and Gordon,D.M. GenBank AY590467.
48. J. Ramon Hernandez, A. Pascual ,L. Martinez-Martinez, 2004, Personal communication.
49. Mendonca, N., E. Ferreira, and M. Canica. 2006. Occurrence of a novel SHV-type enzyme (SHV-55) among isolates of Klebsiella pneumoniae from Portuguese origin in a comparison study for extended-spectrum ß-lactamase-producing evaluation. Diagn. Microbiol. Infect. Dis. 56: 415-420.
50. Dubois,V., Poirel,L., Demarthe,F., Arpin,C., Coulange,L., Minarini,L.A.R., Bezian,M.-C., Nordmann,P. and Quentin,C. 2008. Molecular and biochemical characterization of SHV-56, a novel inhibitor-resistant ß-lactamase from Klebsiella pneumoniae. Antimicrob. Agents Chemother. 52:3792-3794.
51. Ma, L., J. Alba, F. Y. Chang, M. Ishiguro, K. Yamaguchi, L. K. Siu, and Y. Ishii. 2005. Novel SHV-derived extended-spectrum ß-lactamase, SHV-57, that confers resistance to ceftazidime but not cefazolin. Antimicrob Agents Chemother 49:600-605.
52. Lu,Y.M., Zhang,R.Z., Chen,S.W., Wang,S.Y., He,L. and Mu,X.K. GenBank AY790341.
53. Edelstein, M., Ekimov,A. and Stepanova,M. GenBank EU342351.
55. R. Bonomo, 2005, Personal communication.
56. Bao-Dong Ling, Gang Liu,Yong-En Xie, et al. Characterization of a novel extended-spectrum ß-lactamase, SHV-70, from a clinical isolate of Enterobacter cloacae in China. 2006. Int. J. Antimicrob. Agents. 27:355-356.
57. Mendonça, N., E. Ferreira, D. Louro, and M. Caniça. 2009. Molecular epidemiology and antimicrobial susceptibility of extended- and broad-spectrum ß-lactamase-producing Klebsiella pneumoniae isolated in Portugal. Int. J. Antimicrob. Agents 34:29-37.
60. Liu,Z., Wang,H., Yang,Y. and Zuo,B. GenBank DQ174304
61. Liu,Z., Wang,H., Yang,Y. and Zuo,B. GenBank DQ174305
62. Liu,Z., Wang,H., Yang,Y. and Zuo,B. GenBank DQ174306
63. Liu,Z., Wang,H., Yang,Y. and Zuo,B. GenBank DQ174307
64. Liu,Z., Wang,H., Yang,Y. and Zuo,B. GenBank DQ174308
65. Manageiro V., Ferreira E., Caniça M. GenBank AM087453. Biochemical study of a new lactamase inhibitor enzyme (SHV-84) produced by a clinical Escherichia coli strain. Abstract number: P1214. 19th European Congress of Clinical Microbiology and Infectious Diseases. Helsinki, Finland, 16 - 19 May 2009.
65a. Mendonca, N., V. Manageiro, F. Robin, M. J. Salgado, E. Ferreira, M. Canica, and R. Bonnet. 2008. The Lys234Arg substitution in the enzyme SHV-72 is a determinant for resistance to clavulanic acid inhibition. Antimicrob. Agents Chemother. 52:1806-1811.
66. Garcia Dde, O., Y. Doi, D. Szabo, J. M. Adams-Haduch, T. M. Vaz, D. Leite, M. C. Padoveze, M. P. Freire, F. P. Silveira, and D. L. Paterson. 2008. Multiclonal outbreak of Klebsiella pneumoniae producing extended-spectrum ß-lactamase CTX-M-2 and novel variant CTX-M-59 in a neonatal intensive care unit in Brazil. Antimicrob. Agents Chemother. 52:1790-1793.
67. Espinal-Marin,P., Garza-Ramos,U., Reyna-Flores,F. and Silva-Sanchez,J. GenBank DQ328802
68. Curd, E. 2006. Personal communication.
69. Li, J. B., J. Cheng, Q. Wang, Y. Chen, Y. Ye, and X. J. Zhang. 2009. A novel SHV-type ß-lactamase variant (SHV-89) in clinical isolates in China. Mol. Biol. Rep. 36:1141-1148.
70. Machado E, TM Coque, R Cantón, Â Novais, JC Sousa, F Baquero, L Peixe. 2007. High diversity of extended-spectrum ß-lactamases among clinical isolates of Enterobacteriaceae from Portugal. J. Antimicrob. Chemother. 60:1370–1374
71. Lavilla, S., J. J. Gonzalez-Lopez, M. Sabate, A. Garcia-Fernandez, M. N. Larrosa, R. M. Bartolome, A. Carattoli, and G. Prats. 2008. Prevalence of qnr genes among extended-spectrum ß-lactamase-producing enterobacterial isolates in Barcelona, Spain. J. Antimicrob. Chemother. 61:291-295.
72. Sheng,H., Han,L. and Xiao,H. GenBank EF373969.
73. Ramdani-Bouguessa, N., V. Manageiro, D. Jones-Dias, E. Ferreira, M. Tazir, and M. Canica. 2011. Role of SHV ß-lactamase variants in resistance of clinical Klebsiella pneumoniae strains to ß-lactams in an Algerian hospital. J Med Microbiol 60:983-987.
74. Sheng,H., Han,L., Li,H., Zhu,W., Qu,Y., Tian,X., Zhou,J., Zhu,Z., Miao,J., Ni,Y. and Xiao,H. GenBank EF373970.
75. Sheng,H., Han,L., Li,H., Zhu,W., Qu,Y., Tian,X., Zhou,J., Zhu,Z., Miao,J., Ni,Y. and Xiao,H. GenBank EF373972.
76. Sheng,H., Han,L., Li,H., Zhu,W., Qu,Y., Tian,X., Zhou,J., Zhu,Z., Miao,J., Ni,Y. and Xiao,H. GenBank EF373971.
77. Sheng,H., Han,L., Li,H., Zhu,W., Qu,Y., Tian,X., Zhou,J., Zhu,Z., Miao,J., Ni,Y. and Xiao,H. GenBank EF373973. Source reported to be Enterococcus faecalis.
78. Brasme, L., P. Nordmann, F. Fidel, M. F. Lartigue, O. Bajolet, L. Poirel, D. Forte, V. Vernet-Garnier, J. Madoux, J. C. Reveil, C. Alba-Sauviat, I. Baudinat, P. Bineau, C. Bouquigny-Saison, C. Eloy, C. Lafaurie, D. Simeon, J. P. Verquin, F. Noel, C. Strady, and C. De Champs. 2007. Incidence of class A extended-spectrum ß-lactamases in Champagne-Ardenne (France): a 1 year prospective study. J. Antimicrob. Chemother. 60:956-964.
79. Vinue, L., M. Lantero, Y. Saenz, S. Somalo, I. de Diego, F. Perez, F. Ruiz-Larrea, M. Zarazaga, and C. Torres. 2008. Characterization of extended-spectrum ß-lactamases and integrons in Escherichia coli isolates in a Spanish hospital. J. Med. Microbiol. 57:916-920..
80. Abbassi, M. S., C. Torres, W. Achour, L. Vinue, Y. Saenz, D. Costa, O. Bouchami, and A. Ben Hassen. 2008. Genetic characterisation of CTX-M-15-producing Klebsiella pneumoniae and Escherichia coli strains isolated from stem cell transplant patients in Tunisia. Int. J. Antimicrob. Agents. 32:308-314.
81. Ben Achour, N., Belhadj, O., Galleni, M., Ben Moussa, M., and Mercuri, P.S. Study of a natural mutant SHV-type beta -lactamase, SHV-104, from Klebsiella pneumoniae. Int J Microbiol 2014, 548656 (2014)
82. Jones, C. H., A. Ruzin, M. Tuckman, M. A. Visalli, P. J. Petersen, and P. A. Bradford. 2009. Pyrosequencing using the single-nucleotide polymorphism protocol for rapid determination of TEM- and SHV-type extended-spectrum ß-lactamases in clinical isolates and identification of the novel ß-lactamase genes blaSHV-48, blaSHV-105, and blaTEM-155. Antimicrob. Agents Chemother. 53:977-986.
84. Zong, Z., S. R. Partridge, L. Thomas, and J. R. Iredell. 2008. Dominance of blaCTX-M within an Australian extended-spectrum ß-lactamase gene pool. Antimicrob. Agents Chemother. 52:4198-4202.
85. Hammad, A. M., Y. Ishida, and T. Shimamoto. 2009. Prevalence and molecular characterization of ampicillin-resistant Enterobacteriaceae isolated from traditional Egyptian Domiati cheese. J. Food Prot. 72:624-630.
87. Dashti,A.A. and Jadaon,M.M. GenBank EU477409 (partial sequence)
88. Bruderer, T., 2008, personal communication.
89. Paltansing, S., 2008, personal communication
90. Bonomo, R. 2009, personal communication.
91. Doi, Y. 2009, and Zhai,Y., Xu,L. and Gao,Z, 2014, personal communications.
92. Podschun, R., and Pfeifer, Y. GenBank GQ428198.
93. Dashti,A.A., Jadaon,M.M. and Aymes,S.G. GenBank GQ290211.
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95. Lascols, C., M. Hackel, A. M. Hujer, S. H. Marshall, S. K. Bouchillon, D. J. Hoban, S. P. Hawser, R. E. Badal, and R. A. Bonomo. 2012. Using nucleic acid microarrays to perform molecular epidemiology and detect novel ß-lactamases: a snapshot of extended-spectrum ß-lactamases throughout the world. J. Clin. Microbiol. 50:1632-1639.
98. Yamane,K., Hatanaka,K., Matsui,M. and Arakawa,Y. GenBank AB551737.
99. Oteo,J., Sanchez-Romero,I., Fernandez,S., Saez,D. and Campos,J. GenBank HM559945.
100. Jiang,H.-X. and Tang,D. GenBank HQ637576.
101. Wang, P, and Wang,M.G. GenBank HQ661362.
102. Wang. P. and Wang, M.G. GenBank HQ661363.
103. Matasege, L., 2011. Personal communication.
104. Lascols, C., 2011. Personal communication.
105. Bush, K., and Castino. 2011. Personal communication.
106. Bourouis,A., Ben Achour,N., Chihi,H., Belhadj,C., Ben Moussa,M. and Belhadj,O. GenBank GU932590.
106. An, S., J. Chen, Z. Wang, X. Wang, X. Yan, J. Li, Y. Chen, Q. Wang, X. Xu, J. Yang, H. Wang, and Z. Gao. 2012. Predominant characteristics of CTX-M-producing Klebsiella pneumoniae isolates from patients with lower respiratory tract infection in multiple medical centers in China. FEMS Microbiol Lett. 332:137-145.
107. Yan,J.-J., Tsai,L.-H. and Chen,H.-M. GenBank JQ341060
108. Al-Marzooq,F., Yusof,M.Y. and Tay,S.T. GenBank JQ926986.
109. Peixe, L. GenBank JX013655.
110. Piklu Roy Chowdhury, 5/9/12. Personal communication and GenBank KP868752.
111. Castanheira, M., S.E. Farrell, L. M. Deshpande, R.E. Mendes, R.N. Jones. Prevalence of Beta-Lactamase-Encoding Genes among Enterobacteriaceae Bacteremia Isolates Collected in 26 U.S. Hospitals: Report from the SENTRY Antimicrobial Surveillance Program (2010). Antimicrob. Agents Chemother. 2013, 57:3012-3020.
112. Canica, M. 5/24/12. Personal communication.
113. Hansen, Frank. 6/6/12. Personal communication.
114. Dutta, S., S. Mitra, R. Viswanathan, A Saha, and S. Basu. Characterization of novel plasmid-mediated beta-lactamases (SHV-167) and ACT-16 associated with New Delhi Metallo-beta-lactamase 1-harboring isolates from neonates in India. J. Med. Microbiol. 63: 480-482, 2014.
115. AJ Denisuik, PJ Simner, HJ Adam, DJ Hoban and GG Zhanel. GenBank JX870080.
116. Nicole Stoesser. 1/22/13. Personal communication.
117. Hickman, S. 3/13/13. Personal communication & GenBank KF513177.
118. Hickman, S. 5/7/13. Personal communication & GenBank KF513178.
119. Wareham, D. 5/14/13. Personal communication.
120. Gonzalez, J. 6/12./13. Personal communication.
121. Hickman, S. 8/14/13. Personal communication & GenBank KF705209.
122. Hickman, S. 9/24/13. Personal communication & GenBank entries as shown.
123. Izdebski,R., Baraniak,A., Fiett,J. and Gniadkowski,M. Personal communication & GenBank HG934764.
124. Brasme L., Touati A. and De Champs C. 4/29/14. Personal communication.
125. Sridhar Rao, 6/4/14, personal communication and GenBank KM233164 and KM233165
126. Bathoorn, Erik, 7/17/14, personal communication and GenBank LN515533 and LN515534.
127. Rabine, S., 8/5/14, personal communication and GenBank KP050494.
128. Zhai Yao, Yatao Guo, Zhao Zhang, Daixi Li, Yusheng Chen, Jing Wang and Zhancheng Gao. Personal communication & GenBank entries as shown.
129. Ruiz, Belen, 4/17/15, personal communication.

OXA References

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