337:(ADCs) are composed of an antibody linked to a drug. Sortase can be used as a method to link these two molecules. Due to the site-specific ligation of sortase, it shows promise in being used as a method to create ADCs. Sortase poses a potential solution to the challenge of creating homogeneous ADCs where the drug is attached to a single specific site.
29:
383:
bacteria, which recognizes an LPXTG binding motif. Different sortase enzymes found in staphylococcus and other bacteria have other recognition sequences. SrtB for example recognizes a NPQTN binding sequence. These other sortase variants have different properties including different binding motifs and
1521:
Kankainen M, Paulin L, Tynkkynen S, von
Ossowski I, Reunanen J, Partanen P, Satokari R, Vesterlund S, Hendrickx AP, Lebeer S, De Keersmaecker SC, Vanderleyden J, Hämäläinen T, Laukkanen S, Salovuori N, Ritari J, Alatalo E, Korpela R, Mattila-Sandholm T, Lassig A, Hatakka K, Kinnunen KT, Karjalainen
370:
The protein cleaving and ligating function of the sortase enzyme is reliant on the structure of the enzyme binding site and the presence of the correct binding site on the target protein. The requirement of a binding motif limits the versatility of the sortase enzyme and requires the addition of a
404:
The transpeptidase activity of sortase is taken advantage of by structural biologists to produce fusion proteins in vitro. The recognition motif (LPXTG) is added to the C-terminus of a protein of interest while an oligo-glycine motif is added to the N-terminus of the second protein to be ligated.
293:
mutants lacking the srtA gene fail to anchor and display some surface proteins and are impaired in the ability to cause animal infections. Sortase acts on surface proteins that are initiated into the secretion (Sec) pathway and have their signal peptide removed by signal peptidase. The
392:. This process induces mutations in the natural enzyme and selects for mutations that result in the desired properties. SrtA variants have been developed with different binding motifs (LPXSG and LAXTG). Another sortase variant, eSrtA, was specifically developed to have improved
269:
sortase is a transpeptidase that attaches surface proteins to the cell wall; it cleaves between the Gly and Thr of the LPXTG motif and catalyses the formation of an amide bond between the carboxyl-group of threonine and the amino-group of the cell-wall peptidoglycan.
247:), where cell wall LPXTG-mediated decoration has not been reported. Although sortase A, the "housekeeping" sortase, typically acts on many protein targets, other forms of sortase recognize variant forms of the cleavage motif, or catalyze the assembly of
223:. Cleavage occurs between the Thr and Gly, with transient attachment through the Thr residue to the active site Cys residue, followed by transpeptidation that attaches the protein covalently to cell wall components. Sortases occur in almost all
387:
To use the sortase enzyme in broader applications new variations of the enzyme have been developed to exhibit desired properties. SrtA variants that exhibit similar kinetics and catalytic efficiency to the wild type have been engineered using
1575:
Neiers F, Madhurantakam C, Fälker S, Manzano C, Dessen A, Normark S, Henriques-Normark B, Achour A (October 2009). "Two crystal structures of pneumococcal pilus sortase C provide novel insights into catalysis and substrate specificity".
278:
Substrate proteins attached to cell walls by sortases include enzymes, pilins, and adhesion-mediating large surface glycoproteins. These proteins often play important roles in virulence, infection, and colonization by pathogens.
2341:
Bagnoli F, Moschioni M, Donati C, Dimitrovska V, Ferlenghi I, Facciotti C, Muzzi A, Giusti F, Emolo C, Sinisi A, Hilleringmann M, Pansegrau W, Censini S, Rappuoli R, Covacci A, Masignani V, Barocchi MA (August 2008).
340:
A study showed that sortase derived ADCs can effectively kill tumors both in vitro and in vivo. Using sortase to manufacture ADCs may be able to simplify the production and reduce materials needed for the process.
1904:
Solovyova AS, Pointon JA, Race PR, Smith WD, Kehoe MA, Banfield MJ (February 2010). "Solution structure of the major (Spy0128) and minor (Spy0125 and Spy0130) pili subunits from
Streptococcus pyogenes".
282:
Surface proteins not only promote interaction between the invading pathogen and animal tissues, but also provide ingenious strategies for bacterial escape from the host's immune response. In the case of
326:
The sortases are thought to be good targets for new antibiotics as they are important proteins for pathogenic bacteria and some limited commercial interest has been noted by at least one company.
348:
of the natural enzyme. Using error prone PCR to generate mutants of SrtA, the most commonly used natural sortase variant, has been successful in generating more efficient sortase variants.
1824:
Schlüter S, Franz CM, Gesellchen F, Bertinetti O, Herberg FW, Schmidt FR (August 2009). "The high biofilm-encoding Bee locus: a second pilus gene cluster in
Enterococcus faecalis?".
1607:"A collagen-binding adhesin, Acb, and ten other putative MSCRAMM and pilus family proteins of Streptococcus gallolyticus subsp. gallolyticus (Streptococcus bovis Group, biotype I)"
2549:
Manetti AG, Zingaretti C, Falugi F, Capo S, Bombaci M, Bagnoli F, Gambellini G, Bensi G, Mora M, Edwards AM, Musser JM, Graviss EA, Telford JL, Grandi G, Margarit I (May 2007).
983:
Remy
Gebleux, Manfred Briendl, Ulf Grawunder, Roger R Beerli (June 4, 2019). "Sortase a Enzyme-Mediated Generation of Site-Specifically Conjugated Antibody–Drug Conjugates".
1987:
Kang HJ, Middleditch M, Proft T, Baker EN (December 2009). "Isopeptide bonds in bacterial pili and their characterization by X-ray crystallography and mass spectrometry".
2799:
Dramsi S, Caliot E, Bonne I, Guadagnini S, Prévost MC, Kojadinovic M, Lalioui L, Poyart C, Trieu-Cuot P (June 2006). "Assembly and role of pili in group B streptococci".
1307:
Kobashigawa Y, Kumeta H, Ogura K, Inagaki F (March 2009). "Attachment of an NMR-invisible solubility enhancement tag using a sortase-mediated protein ligation method".
2508:"Relative contributions of Enterococcus faecalis OG1RF sortase-encoding genes, srtA and bps (srtC), to biofilm formation and a murine model of urinary tract infection"
169:
113:
101:
1605:
Sillanpää J, Nallapareddy SR, Qin X, Singh KV, Muzny DM, Kovar CL, Nazareth LV, Gibbs RA, Ferraro MJ, Steckelberg JM, Weinstock GM, Murray BE (November 2009).
2764:
Rosini R, Rinaudo CD, Soriani M, Lauer P, Mora M, Maione D, Taddei A, Santi I, Ghezzo C, Brettoni C, Buccato S, Margarit I, Grandi G, Telford JL (July 2006).
771:
Mazmanian SK, Liu G, Ton-That H, Schneewind O (July 1999). "Staphylococcus aureus sortase, an enzyme that anchors surface proteins to the cell wall".
405:
Upon addition of sortase to the protein mixture, the two peptides are covalently linked through a native peptide bond. This reaction is employed by
962:
516:
Mazmanian SK, Ton-That H, Schneewind O (June 2001). "Sortase-catalysed anchoring of surface proteins to the cell wall of
Staphylococcus aureus".
1736:"The structure of the Staphylococcus aureus sortase-substrate complex reveals how the universally conserved LPXTG sorting signal is recognized"
215:. For most substrates of sortase enzymes, the recognition signal consists of the motif LPXTG (Leu-Pro-any-Thr-Gly), then a highly hydrophobic
1650:"Expression, purification, crystallization and preliminary crystallographic analysis of SpaA, a major pilin from Corynebacterium diphtheriae"
1000:
81:
1417:"Structure analysis and site-directed mutagenesis of defined key residues and motives for pilus-related sortase C1 in group B Streptococcus"
459:"Structure analysis and site-directed mutagenesis of defined key residues and motives for pilus-related sortase C1 in group B Streptococcus"
1470:"The Corynebacterium diphtheriae shaft pilin SpaA is built of tandem Ig-like modules with stabilizing isopeptide and disulfide bonds"
1026:"Sortase Enzyme-Mediated Generation of Site-Specifically Conjugated Antibody Drug Conjugates with High In Vitro and In Vivo Potency"
1781:"Intramolecular isopeptide bonds give thermodynamic and proteolytic stability to the major pilin protein of Streptococcus pyogenes"
1415:
Cozzi R, Malito E, Nuccitelli A, D'Onofrio M, Martinelli M, Ferlenghi I, Grandi G, Telford JL, Maione D, Rinaudo CD (June 2011).
457:
Cozzi R, Malito E, Nuccitelli A, D'Onofrio M, Martinelli M, Ferlenghi I, Grandi G, Telford JL, Maione D, Rinaudo CD (June 2011).
367:
Another sub-family of sortases (C60B in MEROPS) contains bacterial sortase B proteins that are approximately 200 residues long.
289:
protein A, immunoglobulins are captured on the microbial surface and camouflage bacteria during the invasion of host tissues.
867:"Archaeosortases and exosortases are widely distributed systems linking membrane transit with posttranslational modification"
2024:
Manzano C, Contreras-Martel C, El
Mortaji L, Izoré T, Fenel D, Vernet T, Schoehn G, Di Guilmi AM, Dessen A (December 2008).
189:
2202:
Fälker S, Nelson AL, Morfeldt E, Jonas K, Hultenby K, Ries J, Melefors O, Normark S, Henriques-Normark B (November 2008).
2344:"A second pilus type in Streptococcus pneumoniae is prevalent in emerging serotypes and mediates adhesion to host cells"
2300:"Sortase A utilizes an ancillary protein anchor for efficient cell wall anchoring of pili in Streptococcus agalactiae"
409:
spectroscopists to produce NMR invisible solubility tags and by X-ray crystallographers to promote complex formation.
3042:
1524:"Comparative genomic analysis of Lactobacillus rhamnosus GG reveals pili containing a human- mucus binding protein"
1352:"Structural basis for activation and non-canonical catalysis of the Rap GTPase activating protein domain of plexin"
1140:
Pallen MJ, Lam AC, Antonio M, Dunbar K (March 2001). "An embarrassment of sortases - a richness of substrates?".
125:
2766:"Identification of novel genomic islands coding for antigenic pilus-like structures in Streptococcus agalactiae"
958:
2159:"The molecular switch that activates the cell wall anchoring step of pilus assembly in gram-positive bacteria"
177:
334:
233:
2430:"Housekeeping sortase facilitates the cell wall anchoring of pilus polymers in Corynebacterium diphtheriae"
525:
224:
1085:"Improved variants of SrtA for site-specific conjugation on antibodies and proteins with high efficiency"
380:
285:
265:
2594:"Corynebacterium diphtheriae employs specific minor pilins to target human pharyngeal epithelial cells"
173:
2723:"Type III pilus of corynebacteria: Pilus length is determined by the level of its major pilin subunit"
1691:
Guttilla IK, Gaspar AH, Swierczynski A, Swaminathan A, Dwivedi P, Das A, Ton-That H (September 2009).
302:
has evolved more than one pathway for the transport of 20 surface proteins to the cell wall envelope.
2258:
1535:
1481:
1253:
1096:
1037:
819:
725:
617:
216:
94:
1734:
Suree N, Liew CK, Villareal VA, Thieu W, Fadeev EA, Clemens JJ, Jung ME, Clubb RT (September 2009).
530:
130:
2635:
Nallapareddy SR, Singh KV, Sillanpää J, Garsin DA, Höök M, Erlandsen SL, Murray BE (October 2006).
1182:"Challenges in the use of sortase and other peptide ligases for site-specific protein modification"
714:"Crystal structure of Spy0129, a Streptococcus pyogenes class B sortase involved in pilus assembly"
2061:"Pili in Gram-negative and Gram-positive bacteria - structure, assembly and their role in disease"
2824:
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2012:
1930:
1892:
1849:
1456:
1332:
939:
551:
498:
389:
357:
379:
The most widely used sortase in biological and medical applications is the SrtA enzyme found in
371:
short protein tag in cases when the desired protein doesn’t contain the necessary binding site.
1522:
H, Saxelin M, Laakso K, Surakka A, Palva A, Salusjärvi T, Auvinen P, de Vos WM (October 2009).
569:
Pallen MJ, Chaudhuri RR, Henderson IR (October 2003). "Genomic analysis of secretion systems".
3001:
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2145:
2092:
2047:
2004:
1975:
1922:
1884:
1841:
1812:
1767:
1722:
1693:"Acyl enzyme intermediates in sortase-catalyzed pilus morphogenesis in gram-positive bacteria"
1679:
1636:
1593:
1563:
1509:
1448:
1405:
1383:
1324:
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2527:
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2319:
2311:
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2180:
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1996:
1965:
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1712:
1704:
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1626:
1618:
1585:
1553:
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1499:
1489:
1438:
1428:
1373:
1363:
1316:
1279:
1261:
1209:
1193:
1149:
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988:
923:
886:
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837:
827:
780:
743:
733:
684:
676:
635:
625:
578:
535:
480:
470:
438:
243:
364:
peptidase family C60 (clan C-) and include the members of several subfamilies of sortases.
156:
1240:
Dorr, Brent M.; Ham, Hyun Ok; An, Chihui; Chaikof, Elliot L.; Liu, David R. (2014-09-16).
2262:
2087:
2060:
1654:
Acta
Crystallographica. Section F, Structural Biology and Crystallization Communications
1539:
1485:
1257:
1100:
1041:
914:
Maresso AW, Schneewind O (March 2008). "Sortase as a target of anti-infective therapy".
823:
729:
621:
2916:"Sortases and pilin elements involved in pilus assembly of Corynebacterium diphtheriae"
2862:
2837:
2747:
2722:
2661:
2636:
2618:
2593:
2532:
2507:
2454:
2429:
2428:
Swaminathan A, Mandlik A, Swierczynski A, Gaspar A, Das A, Ton-That H (November 2007).
2411:
2386:
2368:
2343:
2324:
2299:
2281:
2246:
2228:
2203:
2185:
2158:
2140:
2113:
1970:
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1807:
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1717:
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1504:
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1351:
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1214:
1117:
1084:
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1025:
891:
866:
748:
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689:
664:
640:
605:
423:
310:
106:
2245:
Budzik JM, Marraffini LA, Souda P, Whitelegge JP, Faull KF, Schneewind O (July 2008).
1944:"Sortase D forms the covalent bond that links BcpB to the tip of Bacillus cereus pili"
1153:
3031:
2996:
2979:
2932:
2915:
2838:"Assembly of distinct pilus structures on the surface of Corynebacterium diphtheriae"
2812:
2782:
2765:
2696:
2679:
2609:
2567:
2550:
2489:
2472:
2445:
2219:
1879:
1862:
1336:
842:
807:
539:
228:
86:
2853:
2828:
2713:
2584:
2551:"Streptococcus pyogenes pili promote pharyngeal cell adhesion and biofilm formation"
1934:
1896:
1853:
1460:
808:"Sortase, a universal target for therapeutic agents against gram-positive bacteria?"
555:
502:
62:
2949:
Ton-That H, Schneewind O (May 2004). "Assembly of pili in Gram-positive bacteria".
2016:
152:
2104:
943:
396:, while still other variants were developed to operate in the absence of calcium.
118:
3023:
2897:
2880:
1050:
784:
738:
665:"Roles of the sortases of Streptococcus pneumoniae in assembly of the RlrA pilus"
74:
992:
418:
2251:
Proceedings of the
National Academy of Sciences of the United States of America
2163:
Proceedings of the
National Academy of Sciences of the United States of America
1528:
Proceedings of the
National Academy of Sciences of the United States of America
1474:
Proceedings of the National Academy of Sciences of the United States of America
812:
Proceedings of the National Academy of Sciences of the United States of America
610:
Proceedings of the National Academy of Sciences of the United States of America
298:
genome encodes two sets of sortase and secretion genes. It is conceivable that
2962:
2298:
Nobbs AH, Rosini R, Rinaudo CD, Maione D, Grandi G, Telford JL (August 2008).
2204:"Sortase-mediated assembly and surface topology of adhesive pneumococcal pili"
2077:
2042:
2025:
1918:
1837:
1665:
1589:
1320:
582:
428:
306:
205:
1275:
1205:
211:
that modify surface proteins by recognizing and cleaving a carboxyl-terminal
2387:"SipA is required for pilus formation in Streptococcus pyogenes serotype M3"
2271:
2175:
1797:
1752:
1548:
1494:
1266:
832:
630:
35:
3005:
2970:
2941:
2906:
2871:
2820:
2791:
2756:
2705:
2670:
2627:
2576:
2541:
2498:
2463:
2420:
2377:
2333:
2290:
2237:
2194:
2149:
2130:
2096:
2051:
2008:
1979:
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1926:
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1845:
1816:
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1683:
1640:
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1567:
1513:
1452:
1387:
1328:
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1223:
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935:
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792:
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649:
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547:
494:
90:
3019:
2523:
2315:
1433:
1416:
927:
712:
Kang HJ, Coulibaly F, Proft T, Baker EN (January 2011). Hofmann A (ed.).
475:
458:
220:
69:
2738:
2680:"Pili with strong attachments: Gram-positive bacteria do it differently"
2402:
2359:
1708:
1622:
1443:
1368:
1181:
882:
680:
485:
313:
are functionally analogous, while not in any way homologous to sortase.
1197:
433:
238:
2000:
1468:
Kang HJ, Paterson NG, Gaspar AH, Ton-That H, Baker EN (October 2009).
1409:
1108:
2881:"Protein sorting to the cell wall envelope of Gram-positive bacteria"
2652:
2473:"Assembly of pili on the surface of Bacillus cereus vegetative cells"
2026:"Sortase-mediated pilus fiber biogenesis in Streptococcus pneumoniae"
1863:"Linkage of T3 and Cpa pilins in the Streptococcus pyogenes M3 pilus"
361:
208:
184:
1861:
Quigley BR, Zähner D, Hatkoff M, Thanassi DG, Scott JR (June 2009).
2637:"Endocarditis and biofilm-associated pili of Enterococcus faecalis"
28:
252:
248:
3015:
2980:"Assembly of pili on the surface of Corynebacterium diphtheriae"
146:
57:
2506:
Kemp KD, Singh KV, Nallapareddy SR, Murray BE (November 2007).
2114:"Cell wall anchor structure of BcpA pili in Bacillus anthracis"
1350:
Wang Y, Pascoe HG, Brautigam CA, He H, Zhang X (October 2013).
1180:
Morgan, Holly E.; Turnbull, W. Bruce; Webb, Michael E. (2022).
987:. Methods in Molecular Biology. Vol. 2012. pp. 1–13.
344:
A challenge with using sortase for ADC preparation is the poor
406:
2885:
Biochimica et Biophysica Acta (BBA) - Molecular Cell Research
33:
Pilus-related Sortase C of Group B Streptococcus. PDB entry
2592:
Mandlik A, Swierczynski A, Das A, Ton-That H (April 2007).
1024:
Beerli RR, Hell T, Merkel AS, Grawunder U (July 1, 2015).
2879:
Ton-That H, Marraffini LA, Schneewind O (November 2004).
2471:
Budzik JM, Marraffini LA, Schneewind O (October 2007).
3014:
This article incorporates text from the public domain
2914:
Ton-That H, Marraffini LA, Schneewind O (July 2004).
2247:"Amide bonds assemble pili on the surface of bacilli"
183:
163:
145:
140:
124:
112:
100:
80:
68:
56:
48:
43:
21:
1242:"Reprogramming the specificity of sortase enzymes"
219:, followed by a cluster of basic residues such as
604:Oh SY, Budzik JM, Schneewind O (September 2008).
2112:Budzik JM, Oh SY, Schneewind O (December 2008).
663:LeMieux J, Woody S, Camilli A (September 2008).
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2721:Swierczynski A, Ton-That H (September 2006).
8:
1942:Budzik JM, Oh SY, Schneewind O (May 2009).
606:"Sortases make pili from three ingredients"
2978:Ton-That H, Schneewind O (November 2003).
137:
27:
2995:
2931:
2896:
2861:
2781:
2746:
2695:
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2617:
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2531:
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2410:
2367:
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2184:
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2129:
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2041:
1969:
1959:
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1806:
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1503:
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1432:
1377:
1367:
1283:
1265:
1213:
1116:
1059:
1049:
890:
841:
831:
747:
737:
688:
639:
629:
529:
484:
474:
2836:Gaspar AH, Ton-That H (February 2006).
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963:U.S. Securities and Exchange Commission
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1740:The Journal of Biological Chemistry
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2933:10.1111/j.1365-2958.2004.04117.x
2813:10.1111/j.1365-2958.2006.05190.x
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1880:10.1111/j.1365-2958.2009.06727.x
985:Enzyme-Mediated Ligation Methods
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2854:10.1128/JB.188.4.1526-1533.2006
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571:Current Opinion in Microbiology
1:
1154:10.1016/S0966-842X(01)01956-4
141:Available protein structures:
2898:10.1016/j.bbamcr.2004.04.014
1578:Journal of Molecular Biology
1051:10.1371/journal.pone.0131177
785:10.1126/science.285.5428.760
739:10.1371/journal.pone.0015969
16:Group of prokaryotic enzymes
1907:European Biophysics Journal
1309:Journal of Biomolecular NMR
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3059:
3013:
2963:10.1016/j.tim.2004.03.004
2078:10.1007/s00018-008-8477-4
2043:10.1016/j.str.2008.10.007
1919:10.1007/s00249-009-0432-2
1838:10.1007/s00284-009-9422-y
1666:10.1107/S1744309109027596
1590:10.1016/j.jmb.2009.08.058
1321:10.1007/s10858-008-9296-5
583:10.1016/j.mib.2003.09.005
400:Use in structural biology
317:Pharmaceutic Applications
136:
26:
1186:Chemical Society Reviews
335:Antibody drug conjugates
330:Antibody Drug Conjugates
2842:Journal of Bacteriology
2727:Journal of Bacteriology
2391:Journal of Bacteriology
2348:Journal of Bacteriology
2272:10.1073/pnas.0803565105
2176:10.1073/pnas.0806350105
1798:10.1074/jbc.M109.014514
1753:10.1074/jbc.M109.022624
1697:Journal of Bacteriology
1611:Journal of Bacteriology
1549:10.1073/pnas.0908876106
1495:10.1073/pnas.0906826106
1267:10.1073/pnas.1411179111
916:Pharmacological Reviews
871:Journal of Bacteriology
833:10.1073/pnas.97.10.5013
669:Journal of Bacteriology
631:10.1073/pnas.0807334105
384:reaction efficiencies.
322:As an antibiotic target
234:Shewanella putrefaciens
2984:Molecular Microbiology
2951:Trends in Microbiology
2920:Molecular Microbiology
2801:Molecular Microbiology
2770:Molecular Microbiology
2684:Molecular Microbiology
2598:Molecular Microbiology
2555:Molecular Microbiology
2512:Infection and Immunity
2477:Molecular Microbiology
2434:Molecular Microbiology
2304:Infection and Immunity
2208:Molecular Microbiology
2131:10.1074/jbc.M806796200
1961:10.1074/jbc.M900927200
1867:Molecular Microbiology
1142:Trends in Microbiology
518:Molecular Microbiology
225:Gram-positive bacteria
217:transmembrane sequence
381:staphylococcus aureus
266:Staphylococcus aureus
204:refers to a group of
2524:10.1128/IAI.00663-07
2316:10.1128/IAI.01613-07
1826:Current Microbiology
1434:10.1096/fj.10-174797
928:10.1124/pr.107.07110
476:10.1096/fj.10-174797
2739:10.1128/JB.00606-06
2403:10.1128/JB.01520-07
2360:10.1128/JB.00384-08
2263:2008PNAS..10510215B
1709:10.1128/JB.00627-09
1623:10.1128/JB.00909-09
1540:2009PNAS..10617193K
1486:2009PNAS..10616967K
1369:10.7554/eLife.01279
1258:2014PNAS..11113343D
1252:(37): 13343–13348.
1101:2016NatSR...631899C
1042:2015PLoSO..1031177B
883:10.1128/JB.06026-11
824:2000PNAS...97.5013C
730:2011PLoSO...615969K
681:10.1128/JB.00379-08
622:2008PNAS..10513703O
375:Structural Variants
358:cysteine peptidases
227:and the occasional
1198:10.1039/D0CS01148G
390:directed evolution
3043:Membrane proteins
2001:10.1002/bip.21170
1192:(10): 4121–4145.
1109:10.1038/srep31899
1002:978-1-4939-9545-5
429:SpyTag/SpyCatcher
346:reaction kinetics
199:
198:
195:
194:
190:structure summary
3050:
3009:
2999:
2974:
2945:
2935:
2910:
2900:
2875:
2865:
2832:
2795:
2785:
2760:
2750:
2717:
2699:
2674:
2664:
2653:10.1172/JCI29021
2647:(10): 2799–807.
2631:
2621:
2588:
2570:
2545:
2535:
2518:(11): 5399–404.
2502:
2492:
2467:
2457:
2424:
2414:
2381:
2371:
2337:
2327:
2294:
2284:
2274:
2257:(29): 10215–20.
2241:
2231:
2198:
2188:
2178:
2169:(37): 14147–52.
2153:
2143:
2133:
2124:(52): 36676–86.
2108:
2090:
2080:
2055:
2045:
2020:
1983:
1973:
1963:
1954:(19): 12989–97.
1938:
1900:
1882:
1857:
1820:
1810:
1800:
1791:(31): 20729–37.
1775:
1765:
1755:
1746:(36): 24465–77.
1730:
1720:
1687:
1677:
1644:
1634:
1601:
1571:
1561:
1551:
1517:
1507:
1497:
1480:(40): 16967–71.
1464:
1446:
1436:
1412:
1392:
1391:
1381:
1371:
1347:
1341:
1340:
1304:
1298:
1297:
1287:
1269:
1237:
1228:
1227:
1217:
1177:
1166:
1165:
1137:
1131:
1130:
1120:
1080:
1074:
1073:
1063:
1053:
1021:
1015:
1014:
980:
974:
973:
971:
969:
954:
948:
947:
911:
905:
904:
894:
862:
856:
855:
845:
835:
803:
797:
796:
768:
762:
761:
751:
741:
709:
703:
702:
692:
660:
654:
653:
643:
633:
601:
595:
594:
566:
560:
559:
533:
513:
507:
506:
488:
478:
454:
231:bacterium (e.g.
138:
38:
31:
19:
3058:
3057:
3053:
3052:
3051:
3049:
3048:
3047:
3028:
3027:
3026:
3012:
2977:
2948:
2913:
2891:(1–3): 269–78.
2878:
2835:
2798:
2763:
2733:(17): 6318–25.
2720:
2677:
2634:
2591:
2548:
2505:
2470:
2427:
2384:
2354:(15): 5480–92.
2340:
2297:
2244:
2201:
2156:
2111:
2058:
2036:(12): 1838–48.
2023:
1995:(12): 1126–34.
1986:
1941:
1903:
1860:
1823:
1778:
1733:
1703:(18): 5603–12.
1690:
1660:(Pt 8): 802–4.
1647:
1617:(21): 6643–53.
1604:
1574:
1534:(40): 17193–8.
1520:
1467:
1414:
1404:
1400:
1398:Further reading
1395:
1349:
1348:
1344:
1306:
1305:
1301:
1239:
1238:
1231:
1179:
1178:
1169:
1139:
1138:
1134:
1082:
1081:
1077:
1036:(7): e0131177.
1023:
1022:
1018:
1003:
982:
981:
977:
967:
965:
956:
955:
951:
913:
912:
908:
864:
863:
859:
805:
804:
800:
779:(5428): 760–3.
770:
769:
765:
711:
710:
706:
675:(17): 6002–13.
662:
661:
657:
616:(37): 13703–4.
603:
602:
598:
568:
567:
563:
531:10.1.1.513.4509
515:
514:
510:
456:
455:
451:
447:
415:
402:
377:
354:
332:
324:
319:
276:
274:Biological role
261:
102:OPM superfamily
39:
34:
17:
12:
11:
5:
3056:
3054:
3046:
3045:
3040:
3030:
3029:
3011:
3010:
2990:(4): 1429–38.
2975:
2946:
2911:
2876:
2848:(4): 1526–33.
2833:
2807:(6): 1401–13.
2796:
2761:
2718:
2675:
2632:
2589:
2546:
2503:
2483:(2): 495–510.
2468:
2425:
2382:
2338:
2310:(8): 3550–60.
2295:
2242:
2214:(3): 595–607.
2199:
2154:
2109:
2056:
2021:
1984:
1939:
1901:
1873:(6): 1379–94.
1858:
1821:
1776:
1731:
1688:
1645:
1602:
1572:
1518:
1465:
1427:(6): 1874–86.
1401:
1399:
1396:
1394:
1393:
1342:
1299:
1229:
1167:
1132:
1075:
1016:
1001:
975:
959:"Schedule 14A"
949:
906:
857:
818:(10): 5013–5.
798:
763:
704:
655:
596:
561:
524:(5): 1049–57.
508:
469:(6): 1874–86.
448:
446:
443:
442:
441:
436:
431:
426:
424:Bioengineering
421:
414:
411:
401:
398:
376:
373:
356:This group of
353:
350:
331:
328:
323:
320:
318:
315:
311:archaeosortase
275:
272:
260:
257:
213:sorting signal
197:
196:
193:
192:
187:
181:
180:
167:
161:
160:
150:
143:
142:
134:
133:
128:
122:
121:
116:
110:
109:
104:
98:
97:
84:
78:
77:
72:
66:
65:
60:
54:
53:
50:
46:
45:
41:
40:
32:
24:
23:
22:Sortase family
15:
13:
10:
9:
6:
4:
3:
2:
3055:
3044:
3041:
3039:
3036:
3035:
3033:
3025:
3021:
3017:
3007:
3003:
2998:
2993:
2989:
2985:
2981:
2976:
2972:
2968:
2964:
2960:
2957:(5): 228–34.
2956:
2952:
2947:
2943:
2939:
2934:
2929:
2926:(1): 251–61.
2925:
2921:
2917:
2912:
2908:
2904:
2899:
2894:
2890:
2886:
2882:
2877:
2873:
2869:
2864:
2859:
2855:
2851:
2847:
2843:
2839:
2834:
2830:
2826:
2822:
2818:
2814:
2810:
2806:
2802:
2797:
2793:
2789:
2784:
2779:
2776:(1): 126–41.
2775:
2771:
2767:
2762:
2758:
2754:
2749:
2744:
2740:
2736:
2732:
2728:
2724:
2719:
2715:
2711:
2707:
2703:
2698:
2693:
2690:(2): 320–30.
2689:
2685:
2681:
2676:
2672:
2668:
2663:
2658:
2654:
2650:
2646:
2642:
2638:
2633:
2629:
2625:
2620:
2615:
2611:
2607:
2604:(1): 111–24.
2603:
2599:
2595:
2590:
2586:
2582:
2578:
2574:
2569:
2564:
2561:(4): 968–83.
2560:
2556:
2552:
2547:
2543:
2539:
2534:
2529:
2525:
2521:
2517:
2513:
2509:
2504:
2500:
2496:
2491:
2486:
2482:
2478:
2474:
2469:
2465:
2461:
2456:
2451:
2447:
2443:
2440:(4): 961–74.
2439:
2435:
2431:
2426:
2422:
2418:
2413:
2408:
2404:
2400:
2397:(2): 527–35.
2396:
2392:
2388:
2383:
2379:
2375:
2370:
2365:
2361:
2357:
2353:
2349:
2345:
2339:
2335:
2331:
2326:
2321:
2317:
2313:
2309:
2305:
2301:
2296:
2292:
2288:
2283:
2278:
2273:
2268:
2264:
2260:
2256:
2252:
2248:
2243:
2239:
2235:
2230:
2225:
2221:
2217:
2213:
2209:
2205:
2200:
2196:
2192:
2187:
2182:
2177:
2172:
2168:
2164:
2160:
2155:
2151:
2147:
2142:
2137:
2132:
2127:
2123:
2119:
2115:
2110:
2106:
2102:
2098:
2094:
2089:
2084:
2079:
2074:
2071:(4): 613–35.
2070:
2066:
2062:
2057:
2053:
2049:
2044:
2039:
2035:
2031:
2027:
2022:
2018:
2014:
2010:
2006:
2002:
1998:
1994:
1990:
1985:
1981:
1977:
1972:
1967:
1962:
1957:
1953:
1949:
1945:
1940:
1936:
1932:
1928:
1924:
1920:
1916:
1913:(3): 469–80.
1912:
1908:
1902:
1898:
1894:
1890:
1886:
1881:
1876:
1872:
1868:
1864:
1859:
1855:
1851:
1847:
1843:
1839:
1835:
1832:(2): 206–11.
1831:
1827:
1822:
1818:
1814:
1809:
1804:
1799:
1794:
1790:
1786:
1782:
1777:
1773:
1769:
1764:
1759:
1754:
1749:
1745:
1741:
1737:
1732:
1728:
1724:
1719:
1714:
1710:
1706:
1702:
1698:
1694:
1689:
1685:
1681:
1676:
1671:
1667:
1663:
1659:
1655:
1651:
1646:
1642:
1638:
1633:
1628:
1624:
1620:
1616:
1612:
1608:
1603:
1599:
1595:
1591:
1587:
1584:(3): 704–16.
1583:
1579:
1573:
1569:
1565:
1560:
1555:
1550:
1545:
1541:
1537:
1533:
1529:
1525:
1519:
1515:
1511:
1506:
1501:
1496:
1491:
1487:
1483:
1479:
1475:
1471:
1466:
1462:
1458:
1454:
1450:
1445:
1440:
1435:
1430:
1426:
1422:
1421:FASEB Journal
1418:
1411:
1407:
1403:
1402:
1397:
1389:
1385:
1380:
1375:
1370:
1365:
1361:
1357:
1353:
1346:
1343:
1338:
1334:
1330:
1326:
1322:
1318:
1315:(3): 145–50.
1314:
1310:
1303:
1300:
1295:
1291:
1286:
1281:
1277:
1273:
1268:
1263:
1259:
1255:
1251:
1247:
1243:
1236:
1234:
1230:
1225:
1221:
1216:
1211:
1207:
1203:
1199:
1195:
1191:
1187:
1183:
1176:
1174:
1172:
1168:
1163:
1159:
1155:
1151:
1148:(3): 97–102.
1147:
1143:
1136:
1133:
1128:
1124:
1119:
1114:
1110:
1106:
1102:
1098:
1094:
1090:
1086:
1079:
1076:
1071:
1067:
1062:
1057:
1052:
1047:
1043:
1039:
1035:
1031:
1027:
1020:
1017:
1012:
1008:
1004:
998:
994:
990:
986:
979:
976:
964:
960:
953:
950:
945:
941:
937:
933:
929:
925:
922:(1): 128–41.
921:
917:
910:
907:
902:
898:
893:
888:
884:
880:
876:
872:
868:
861:
858:
853:
849:
844:
839:
834:
829:
825:
821:
817:
813:
809:
802:
799:
794:
790:
786:
782:
778:
774:
767:
764:
759:
755:
750:
745:
740:
735:
731:
727:
724:(1): e15969.
723:
719:
715:
708:
705:
700:
696:
691:
686:
682:
678:
674:
670:
666:
659:
656:
651:
647:
642:
637:
632:
627:
623:
619:
615:
611:
607:
600:
597:
592:
588:
584:
580:
577:(5): 519–27.
576:
572:
565:
562:
557:
553:
549:
545:
541:
537:
532:
527:
523:
519:
512:
509:
504:
500:
496:
492:
487:
482:
477:
472:
468:
464:
463:FASEB Journal
460:
453:
450:
444:
440:
437:
435:
432:
430:
427:
425:
422:
420:
417:
416:
412:
410:
408:
399:
397:
395:
391:
385:
382:
374:
372:
368:
365:
363:
359:
351:
349:
347:
342:
338:
336:
329:
327:
321:
316:
314:
312:
308:
303:
301:
297:
292:
288:
287:
280:
273:
271:
268:
267:
258:
256:
254:
250:
246:
245:
240:
236:
235:
230:
229:Gram-negative
226:
222:
218:
214:
210:
207:
203:
191:
188:
186:
182:
179:
175:
171:
168:
166:
162:
158:
154:
151:
148:
144:
139:
135:
132:
129:
127:
123:
120:
117:
115:
111:
108:
105:
103:
99:
96:
92:
88:
85:
83:
79:
76:
73:
71:
67:
64:
61:
59:
55:
51:
47:
42:
37:
30:
25:
20:
2987:
2983:
2954:
2950:
2923:
2919:
2888:
2884:
2845:
2841:
2804:
2800:
2773:
2769:
2730:
2726:
2687:
2683:
2644:
2640:
2601:
2597:
2558:
2554:
2515:
2511:
2480:
2476:
2437:
2433:
2394:
2390:
2351:
2347:
2307:
2303:
2254:
2250:
2211:
2207:
2166:
2162:
2121:
2117:
2068:
2064:
2033:
2029:
1992:
1988:
1951:
1947:
1910:
1906:
1870:
1866:
1829:
1825:
1788:
1784:
1743:
1739:
1700:
1696:
1657:
1653:
1614:
1610:
1581:
1577:
1531:
1527:
1477:
1473:
1444:11562/349253
1424:
1420:
1359:
1355:
1345:
1312:
1308:
1302:
1249:
1245:
1189:
1185:
1145:
1141:
1135:
1095:(1): 31899.
1092:
1088:
1078:
1033:
1029:
1019:
984:
978:
966:. Retrieved
952:
919:
915:
909:
877:(1): 36–48.
874:
870:
860:
815:
811:
801:
776:
772:
766:
721:
717:
707:
672:
668:
658:
613:
609:
599:
574:
570:
564:
521:
517:
511:
486:11562/349253
466:
462:
452:
403:
386:
378:
369:
366:
355:
343:
339:
333:
325:
304:
299:
295:
290:
284:
281:
277:
264:
262:
242:
232:
201:
200:
1989:Biopolymers
419:Protein tag
206:prokaryotic
114:OPM protein
44:Identifiers
3032:Categories
1362:: e01279.
968:29 October
445:References
360:belong to
307:exosortase
305:Note that
153:structures
3024:IPR005754
2030:Structure
1413:;
1337:207183676
1276:0027-8424
1206:0306-0012
526:CiteSeerX
352:Structure
300:S. aureus
296:S. aureus
291:S. aureus
286:S. aureus
75:IPR005754
3020:InterPro
3006:14622427
2971:15120142
2942:15225319
2907:15546671
2872:16452436
2829:40698153
2821:16796677
2792:16824100
2757:16923899
2714:25384605
2706:16978260
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