645:. After protein concentration is determined, equal amounts of tissue protein are loaded into a gel. The protein is then allowed to migrate through the gel. After electrophoresis, the gel is put into a renaturing buffer in order to return the cathepsins to their native conformation. The gel is then put into an activation buffer of a specific pH and left to incubate overnight at 37 °C. This activation step allows the cathepsins to degrade the gelatin substrate. When the gel is stained using a
694:, and cathepsin itself. Initial efforts to purify and characterize proteases using hemoglobin transpired at a time when the word "cathepsin" indicated a single enzyme; the existence of multiple, distinct cathepsin family members (e.g. B, H, L) did not appear to be understood at the time. However, by 1937 Bergmann and colleagues began to differentiate cathepsins on the basis of their source in the human body (e.g. liver cathepsin, spleen cathepsin).
1662:
550:, the major component of the non-mineral protein matrix of the bone. Cathepsin K, among other cathepsins, plays a role in cancer metastasis through the degradation of the extracellular matrix. The genetic knockout for cathepsin S and K in mice with atherosclerosis was shown to reduce the size of atherosclerotic lesions. The expression of cathepsin K in cultured endothelial cells is regulated by
3202:
29:
654:
have optimum proteolytic activity. Cathepsin K is able to degrade gelatin at pH 7 and 8, but these pH levels do not allow for cathepsins L and V activity. At a pH 4 cathepsin V is active, but cathepsin K is not. Adjusting the pH of the activation buffer can allow for further identification of cathepsin types.
653:
protocol has been used to detect femtomole quantities of mature cathepsin K. The different cathepsins can be identified based on their migration distance due to their molecular weights: cathepsin K (~37 kDa), V (~35 kDa), S (~25kDa), and L (~20 kDa). Cathepsins have specific pH levels at which they
235:
that degrade proteins) found in all animals as well as other organisms. There are approximately a dozen members of this family, which are distinguished by their structure, catalytic mechanism, and which proteins they cleave. Most of the members become activated at the low pH found in
598:. Cathepsin K inhibitors, Relacatib, Balicatib, and Odanacatib, were terminated during clinical trials at phases I, II, and III, respectively, owing to adverse side effects. SAR114137, a Cathepsin S inhibitor, did not progress past phase I for chronic pain. In 2022,
666:
and Eugen Bamann to describe a proteolytic activity of leukocytes and tissues at slightly acidic pH (Willstätter & Bamann (1929) Hoppe-Seylers Z. Physiol. Chemie 180, 127-143). The earliest record of "cathepsin" found in the
1413:
Platt MO, Ankeny RF, Shi GP, Weiss D, Vega JD, Taylor WR, Jo H (March 2007). "Expression of cathepsin K is regulated by shear stress in cultured endothelial cells and is increased in endothelium in human atherosclerosis".
1484:
Brömme D, Li Z, Barnes M, Mehler E (February 1999). "Human cathepsin V functional expression, tissue distribution, electrostatic surface potential, enzymatic characterization, and chromosomal localization".
1371:
Lutgens E, Lutgens SP, Faber BC, Heeneman S, Gijbels MM, de
Winther MP, Frederik P, van der Made I, Daugherty A, Sijbers AM, Fisher A, Long CJ, Saftig P, Black D, Daemen MJ, Cleutjens KB (January 2006).
679:
in 1949. However, references within this article indicate that cathepsins were first identified and named around the turn of the 20th century. Much of this earlier work was done in the laboratory of
1123:"The cysteine protease inhibitor, E64d, reduces brain amyloid-β and improves memory deficits in Alzheimer's disease animal models by inhibiting cathepsin B, but not BACE1, β-secretase activity"
394:
and "it attenuates the anti-tumor immune response of decaying chemokines to inhibit the function of dendritic cells". Cathepsins B and L are involved in matrix degradation and cell invasion.
562:
Mouse cathepsin L is homologous to human cathepsin V. Mouse cathepsin L has been shown to play a role in adipogenesis and glucose intolerance in mice. Cathepsin L degrades fibronectin,
526:. Unlike some of the other cathepsins, cathepsin D has some protease activity at neutral pH. High levels of this enzyme in tumor cells seems to be associated with greater invasiveness.
633:
with a substrate in order to detect enzyme activity. Cathepsin zymography separates different cathepsins based on their migration through a polyacrylamide gel co-polymerized with a
2387:
2247:
2452:
177:
1987:
686:
It is notable that research published in the 1930s (primarily by
Bergmann) used the term "catheptic enzymes" to refer to a broad family of proteases that included
1178:
Habibollahi, Peiman; Figueiredo, Jose-Luiz; Heidari, Pedram; Dulak, Austin M; Imamura, Yu; Bass, Adam J.; Ogino, Shuji; Chan, Andrew T; Mahmood, Umar (2012).
3036:
2612:
1690:
Li WA, Barry ZT, Cohen JD, Wilder CL, Deeds RJ, Keegan PM, Platt MO (June 2010). "Detection of femtomole quantities of mature cathepsin K with zymography".
475:
is significantly higher than in primary focus lysates. Cathepsin A increased in muscles moderately affected by muscular dystrophy and denervating diseases.
1180:"Optical Imaging with a Cathepsin B Activated Probe for the Enhanced Detection of Esophageal Adenocarcinoma by Dual Channel Fluorescent Upper GI Endoscopy"
570:(IGF-1R). Cathepsin L-deficient mice were shown to have less adipose tissue, lower serum glucose and insulin levels, more insulin receptor subunits, more
2979:
89:
1520:
Yang M, Zhang Y, Pan J, Sun J, Liu J, Libby P, Sukhova GK, Doria A, Katunuma N, Peroni OD, Guerre-Millo M, Kahn BB, Clement K, Shi GP (August 2007).
1374:"Disruption of the cathepsin K gene reduces atherosclerosis progression and induces plaque fibrosis but accelerates macrophage foam cell formation"
2445:
567:
423:
125:
1727:"Manipulating substrate and pH in zymography protocols selectively distinguishes cathepsins K, L, S, and V activity in cells and tissues"
649:, areas of the gel still containing gelatin appear blue. The areas of the gel where cathepsins were active appear as white bands. This
1980:
1449:
Salminen-Mankonen HJ, Morko J, Vuorio E (February 2007). "Role of cathepsin K in normal joints and in the development of arthritis".
2605:
2438:
1663:""Sorrento Therapeutics Announces the FDA IND Clearance of STI-1558, An Oral M(pro) and Cathepsin L Inhibitor to Treat COVID-19""
1229:"Cathepsin D released by lactating rat mammary epithelial cells is involved in prolactin cleavage under physiological conditions"
447:
676:
1569:
BratkoviÄŤ, et al. (2005). "Affinity selection to papain yields potent peptide inhibitors of cathepsins L, B, H, and K.".
2972:
1973:
197:
3132:
990:"Serine protease cathepsin G regulates adhesion-dependent neutrophil effector functions by modulating integrin clustering"
3192:
2207:
603:
582:
Five cyclic peptides show inhibitory activity towards human cathepsins L, B, H, and K. Several inhibitors have reached
3122:
2763:
2758:
2598:
638:
1965:
1271:
375:
Cathepsins are involved in many physiological processes and have been implicated in a number of human diseases. The
420:, Cathepsin B and to a lesser extent cathepsin L have been found to be necessary for the virus to enter host cells.
3232:
2965:
2632:
2472:
2122:
2107:
2007:
499:
491:
498:. Overexpression of the encoded protein, which is a member of the peptidase C1 family, has been associated with
2629:
2502:
2469:
2004:
1959:
646:
759:
Turk, Vito; Stoka, Veronika; Vasiljeva, Olga; Renko, Miha; Sun, Tao; Turk, Boris; Turk, Dušan (January 2012).
185:
2823:
2397:
2362:
2357:
1233:
502:
and other tumors. Cathepsin B has also been implicated in the progression of various human tumors including
637:
substrate. The electrophoresis takes place in non-reducing conditions, and the enzymes are protected from
3174:
3104:
3061:
2989:
663:
407:
402:
3127:
2352:
2202:
428:
94:
812:
181:
3142:
3069:
2342:
1827:
1042:
650:
138:
3237:
3117:
3001:
2916:
2242:
2232:
2049:
2044:
622:
3227:
2465:
1643:
1312:
970:
892:"Inhibition of cathepsin S produces neuroprotective effects after traumatic brain injury in mice"
741:
472:
443:
376:
2625:
2224:
1925:
1863:"On proteolytic enzymes XV. Regarding the general nature of intracellular proteolytic enzymes"
1843:
1797:
1756:
1707:
1635:
1627:
1586:
1551:
1502:
1466:
1431:
1395:
1353:
1304:
1252:
1209:
1160:
1142:
1103:
1086:
Im E, Kazlauskas A (March 2007). "The role of cathepsins in ocular physiology and pathology".
1068:
1011:
962:
923:
872:
837:
790:
733:
546:
are the bone resorbing cells of the body, and they secrete cathepsin K in order to break down
515:
464:
296:
280:
172:
2430:
3051:
2936:
2828:
2818:
2813:
2808:
2803:
2590:
2517:
2512:
2016:
1915:
1907:
1874:
1835:
1787:
1746:
1738:
1699:
1617:
1578:
1541:
1533:
1494:
1458:
1423:
1385:
1343:
1296:
1286:
1272:"Molecular cloning of human cathepsin O, a novel endoproteinase and homologue of rabbit OC2"
1242:
1199:
1191:
1150:
1134:
1095:
1058:
1050:
1001:
954:
943:"Reconsider Alzheimer's disease by the 'calpain-cathepsin hypothesis'--a perspective review"
913:
903:
864:
827:
780:
772:
723:
563:
487:
2957:
2783:
1950:
164:
3222:
3021:
3010:
2257:
2252:
2024:
2000:
270:
253:
813:"Involvement of cathepsins in the invasion, metastasis and proliferation of cancer cells"
1831:
1818:
Bergmann M, Fruton JS (July 1936). "Regarding the general nature of catheptic enzymes".
1046:
118:
3206:
2540:
1920:
1896:"The estimation of cathepsin with hemoglobin and the partial purification of cathepsin"
1895:
1751:
1726:
1546:
1521:
1204:
1179:
1155:
1122:
1063:
1030:
918:
891:
785:
760:
630:
626:
591:
583:
542:, a disease in which a decrease in bone density causes an increased risk for fracture.
503:
1955:
1879:
1862:
1792:
1775:
1390:
1373:
3216:
3178:
3084:
2402:
2275:
2264:
1996:
1291:
213:
130:
1647:
1316:
974:
958:
57:
3094:
3025:
2875:
2871:
2298:
2192:
2034:
868:
745:
680:
595:
587:
551:
539:
495:
433:
338:
332:
160:
1031:"Endosomal Proteolysis of the Ebola Virus Glycoprotein Is Necessary for Infection"
70:
1006:
989:
776:
82:
3089:
3074:
3041:
2993:
2946:
2941:
2911:
2895:
2890:
2885:
2880:
2866:
2861:
2856:
2851:
2846:
2579:
2574:
2556:
2417:
2412:
2061:
1427:
599:
535:
519:
483:
439:
380:
362:
356:
350:
344:
326:
320:
314:
308:
302:
292:
286:
276:
266:
245:
3201:
1839:
1582:
1462:
1099:
761:"Cysteine cathepsins: From structure, function and regulation to new frontiers"
683:, who spent the first several decades of the century defining these proteases.
28:
2926:
2740:
2709:
2704:
2699:
2694:
2535:
2379:
2321:
2102:
2097:
2077:
1742:
618:
543:
249:
241:
1703:
1631:
1270:
Shi GP, Chapman HA, Bhairi SM, DeLeeuw C, Reddy VY, Weiss SJ (January 1995).
1146:
3155:
2931:
2793:
2788:
2778:
2768:
2735:
2689:
2684:
2679:
2674:
2669:
2664:
2659:
2654:
2649:
2393:
2212:
2092:
2082:
1138:
1054:
642:
412:
237:
1929:
1847:
1801:
1760:
1711:
1639:
1590:
1555:
1506:
1470:
1435:
1399:
1357:
1256:
1213:
1164:
1107:
1072:
1015:
966:
927:
876:
841:
794:
737:
1308:
1300:
908:
728:
711:
134:
3079:
3031:
3015:
2921:
2621:
2492:
2461:
2293:
2288:
2237:
2087:
2072:
2039:
1911:
1622:
1605:
1227:
Lkhider M, Castino R, Bouguyon E, Isidoro C, Ollivier-Bousquet M (2004).
890:
Xu J, Wang H, Ding K, Lu X, Li T, Wang J, Wang C, Wang J (Oct 24, 2013).
691:
607:
547:
451:
228:
77:
1348:
1331:
3150:
2750:
2641:
2347:
2337:
2283:
2197:
2029:
668:
634:
523:
391:
240:. Thus, the activity of this family lies almost entirely within those
232:
106:
101:
1498:
1247:
1228:
1195:
2798:
2730:
2725:
2487:
2374:
2316:
1946:
832:
687:
672:
468:
397:
387:
192:
113:
1537:
942:
2773:
2497:
2383:
2369:
2308:
2182:
2177:
2172:
2167:
2162:
2157:
1522:"Cathepsin L activity controls adipogenesis and glucose tolerance"
988:
Raptis SZ, Shapiro SD, Simmons PM, Cheng AM, Pham CT (June 2005).
571:
417:
1776:"The hydrolysis of nucleoproteins by cathepsins from calf thymus"
855:
Lipton P (October 1999). "Ischemic cell death in brain neurons".
2152:
2147:
2142:
2137:
2132:
2127:
2117:
2112:
154:
64:
52:
2961:
2594:
2434:
1969:
554:. Cathepsin K has also been shown to play a role in arthritis.
256:. Cathepsins have a vital role in mammalian cellular turnover.
1332:"Cysteine cathepsins and the cutting edge of cancer invasion"
765:
Biochimica et
Biophysica Acta (BBA) - Proteins and Proteomics
586:, targeting cathepsins K and S as promising therapeutics for
463:
Deficiencies in this protein are linked to multiple forms of
712:"Papain-like peptidases: structure, function, and evolution"
1725:
Wilder CL, Park KY, Keegan PM, Platt MO (December 2011).
1121:
Hook, Gregory; Hook, Vivian; Kindy, Mark (2011-01-01).
574:(GLUT4) and more fibronectin than wild type controls.
3190:
1949:
online database for peptidases and their inhibitors:
1606:"Merck & Co. drops osteoporosis drug odanacatib"
518:) appears to cleave a variety of substrates such as
3141:
3103:
3060:
3050:
3000:
2904:
2837:
2749:
2718:
2640:
2565:
2549:
2528:
2480:
2330:
2307:
2273:
2223:
2060:
2015:
1571:
191:
171:
153:
148:
124:
112:
100:
88:
76:
63:
51:
43:
38:
21:
710:Novinec, Marko; LenarÄŤiÄŤ, Brigita (1 June 2013).
248:, which works extracellularly after secretion by
1813:
1811:
806:
804:
379:have attracted significant research effort as
2973:
2606:
2446:
1981:
8:
3037:Bactericidal permeability-increasing protein
606:clearance to begin phase I studies to treat
3057:
2980:
2966:
2958:
2613:
2599:
2591:
2453:
2439:
2431:
1988:
1974:
1966:
1685:
1683:
145:
27:
1958:at the U.S. National Library of Medicine
1919:
1878:
1791:
1750:
1621:
1545:
1389:
1347:
1290:
1246:
1203:
1154:
1062:
1005:
917:
907:
831:
784:
727:
662:The term cathepsin was coined in 1929 by
534:Cathepsin K is the most potent mammalian
244:. There are, however, exceptions such as
3197:
702:
2001:serine proteases/serine endopeptidases
1861:Bergmann M, Fruton JS (June 1, 1937).
811:Nomura T, Katunuma N (February 2005).
18:
568:insulin-like growth factor 1 receptor
7:
1774:Maver ME, Greco AE (December 1949).
1330:Gocheva V, Joyce JA (January 2007).
602:, a Cathepsin L inhibitor, received
1867:The Journal of Biological Chemistry
1416:Am. J. Physiol. Heart Circ. Physiol
14:
1900:The Journal of General Physiology
1391:10.1161/CIRCULATIONAHA.105.561449
3200:
448:age-related macular degeneration
1894:Anson, M. L. (September 1936).
1661:Cooley, Brian (July 19, 2022).
959:10.1016/j.pneurobio.2013.02.004
677:Journal of Biological Chemistry
1127:Journal of Alzheimer's Disease
869:10.1152/physrev.1999.79.4.1431
538:. Cathepsin K is involved in
467:. The cathepsin A activity in
1:
3133:Eosinophil-derived neurotoxin
2213:Urinary plasminogen activator
1880:10.1016/S0021-9258(18)74429-3
1793:10.1016/S0021-9258(18)56608-4
1610:Nature Reviews Drug Discovery
1604:Mullard, Asher (2016-10-01).
149:Available protein structures:
2208:Tissue plasminogen activator
1292:10.1016/0014-5793(94)01349-6
1007:10.1016/j.immuni.2005.03.015
777:10.1016/j.bbapap.2011.10.002
3123:Eosinophil cationic protein
1428:10.1152/ajpheart.00954.2006
3254:
1840:10.1126/science.84.2169.89
1667:Sorrento Therapeutics, Inc
1583:10.1016/j.bbrc.2005.05.028
1463:10.2174/138945007779940188
1100:10.1016/j.exer.2006.05.017
438:Several ocular disorders:
365:(or X) (cysteine protease)
341:(or V) (cysteine protease)
3168:
2299:Proteinase 3/Myeloblastin
1743:10.1016/j.abb.2011.09.009
896:Mediators of Inflammation
500:esophageal adenocarcinoma
492:amyloid precursor protein
471:of metastatic lesions of
144:
26:
2503:Signal peptide peptidase
1960:Medical Subject Headings
1704:10.1016/j.ab.2010.02.035
33:Structure of Cathepsin K
1234:Journal of Cell Science
1139:10.3233/JAD-2011-110101
1055:10.1126/science.1110656
3175:platelet alpha-granule
2353:Proprotein convertases
1731:Arch. Biochem. Biophys
403:Traumatic brain injury
3128:Eosinophil peroxidase
2203:Plasminogen activator
947:Progress in Neurology
729:10.1515/bmc-2012-0054
716:BioMolecular Concepts
429:Chronic periodontitis
371:Clinical significance
359:(cysteine proteinase)
311:(cysteine proteinase)
3070:Alkaline phosphatase
3002:Azurophilic granules
2343:Prolyl endopeptidase
1912:10.1085/jgp.20.4.565
1623:10.1038/nrd.2016.207
1241:(Pt 21): 5155–5164.
941:Yamashima T (2013).
671:database (e.g., via
651:cathepsin zymography
647:Coomassie blue stain
614:Cathepsin zymography
223:"boil"; abbreviated
3118:Major basic protein
2917:Cancer procoagulant
2466:aspartate proteases
1832:1936Sci....84...89B
1349:10.4161/cc.6.1.3669
1047:2005Sci...308.1643C
1041:(5728): 1643–1645.
1029:Chandran K (2005).
909:10.1155/2013/187873
664:Richard Willstätter
623:gel electrophoresis
572:glucose transporter
408:Alzheimer's disease
390:, Cathepsin D is a
377:cysteine cathepsins
353:(cysteine protease)
347:(cysteine protease)
335:(cysteine protease)
329:(cysteine protease)
323:(cysteine protease)
305:(aspartyl protease)
289:(cysteine protease)
16:Family of proteases
2626:cysteine proteases
486:may function as a
473:malignant melanoma
444:retinal detachment
3188:
3187:
3164:
3163:
3052:Specific granules
2955:
2954:
2588:
2587:
2428:
2427:
2225:Complement system
2017:Digestive enzymes
1499:10.1021/bi982175f
1451:Curr Drug Targets
1248:10.1242/jcs.01396
1196:10.7150/thno.4088
516:aspartyl protease
465:galactosialidosis
317:(serine protease)
297:aspartyl protease
281:cysteine protease
207:
206:
203:
202:
198:structure summary
3245:
3233:Protein families
3205:
3204:
3196:
3058:
3022:serine proteases
2988:Contents of the
2982:
2975:
2968:
2959:
2937:3C-like protease
2615:
2608:
2601:
2592:
2455:
2448:
2441:
2432:
1990:
1983:
1976:
1967:
1934:
1933:
1923:
1891:
1885:
1884:
1882:
1858:
1852:
1851:
1815:
1806:
1805:
1795:
1771:
1765:
1764:
1754:
1722:
1716:
1715:
1687:
1678:
1677:
1675:
1673:
1658:
1652:
1651:
1625:
1601:
1595:
1594:
1566:
1560:
1559:
1549:
1517:
1511:
1510:
1481:
1475:
1474:
1446:
1440:
1439:
1410:
1404:
1403:
1393:
1368:
1362:
1361:
1351:
1327:
1321:
1320:
1294:
1276:
1267:
1261:
1260:
1250:
1224:
1218:
1217:
1207:
1175:
1169:
1168:
1158:
1118:
1112:
1111:
1083:
1077:
1076:
1066:
1026:
1020:
1019:
1009:
985:
979:
978:
938:
932:
931:
921:
911:
902:(2013): 187873.
887:
881:
880:
852:
846:
845:
835:
833:10.2152/jmi.52.1
817:
808:
799:
798:
788:
756:
750:
749:
731:
707:
564:insulin receptor
514:Cathepsin D (an
488:beta-secretase 1
146:
31:
19:
3253:
3252:
3248:
3247:
3246:
3244:
3243:
3242:
3213:
3212:
3211:
3199:
3191:
3189:
3184:
3160:
3137:
3099:
3046:
3011:Myeloperoxidase
2996:
2986:
2956:
2951:
2900:
2833:
2745:
2714:
2636:
2619:
2589:
2584:
2561:
2545:
2524:
2476:
2459:
2429:
2424:
2326:
2303:
2269:
2219:
2056:
2025:Enteropeptidase
2011:
1994:
1942:
1937:
1893:
1892:
1888:
1860:
1859:
1855:
1826:(2169): 89–90.
1817:
1816:
1809:
1773:
1772:
1768:
1724:
1723:
1719:
1689:
1688:
1681:
1671:
1669:
1660:
1659:
1655:
1603:
1602:
1598:
1568:
1567:
1563:
1538:10.1038/ncb1623
1519:
1518:
1514:
1483:
1482:
1478:
1448:
1447:
1443:
1422:(3): H1479–86.
1412:
1411:
1407:
1370:
1369:
1365:
1329:
1328:
1324:
1274:
1269:
1268:
1264:
1226:
1225:
1221:
1177:
1176:
1172:
1120:
1119:
1115:
1085:
1084:
1080:
1028:
1027:
1023:
987:
986:
982:
940:
939:
935:
889:
888:
884:
863:(4): 1431–568.
854:
853:
849:
815:
810:
809:
802:
758:
757:
753:
709:
708:
704:
700:
660:
616:
584:clinical trials
580:
560:
532:
512:
481:
461:
373:
368:
271:serine protease
262:
254:bone resorption
34:
17:
12:
11:
5:
3251:
3249:
3241:
3240:
3235:
3230:
3225:
3215:
3214:
3210:
3209:
3186:
3185:
3183:
3182:
3169:
3166:
3165:
3162:
3161:
3159:
3158:
3153:
3147:
3145:
3139:
3138:
3136:
3135:
3130:
3125:
3120:
3115:
3109:
3107:
3101:
3100:
3098:
3097:
3092:
3087:
3082:
3077:
3072:
3066:
3064:
3055:
3048:
3047:
3045:
3044:
3039:
3034:
3029:
3018:
3013:
3007:
3005:
2998:
2997:
2987:
2985:
2984:
2977:
2970:
2962:
2953:
2952:
2950:
2949:
2944:
2939:
2934:
2929:
2924:
2919:
2914:
2908:
2906:
2902:
2901:
2899:
2898:
2893:
2888:
2883:
2878:
2869:
2864:
2859:
2854:
2849:
2843:
2841:
2835:
2834:
2832:
2831:
2826:
2821:
2816:
2811:
2806:
2801:
2796:
2791:
2786:
2781:
2776:
2771:
2766:
2761:
2755:
2753:
2747:
2746:
2744:
2743:
2738:
2733:
2728:
2722:
2720:
2716:
2715:
2713:
2712:
2707:
2702:
2697:
2692:
2687:
2682:
2677:
2672:
2667:
2662:
2657:
2652:
2646:
2644:
2638:
2637:
2620:
2618:
2617:
2610:
2603:
2595:
2586:
2585:
2583:
2582:
2577:
2571:
2569:
2563:
2562:
2560:
2559:
2553:
2551:
2547:
2546:
2544:
2543:
2541:HIV-1 protease
2538:
2532:
2530:
2526:
2525:
2523:
2522:
2521:
2520:
2515:
2508:Beta secretase
2505:
2500:
2495:
2490:
2484:
2482:
2478:
2477:
2460:
2458:
2457:
2450:
2443:
2435:
2426:
2425:
2423:
2422:
2421:
2420:
2415:
2405:
2400:
2391:
2377:
2372:
2367:
2366:
2365:
2360:
2350:
2345:
2340:
2334:
2332:
2328:
2327:
2325:
2324:
2319:
2313:
2311:
2305:
2304:
2302:
2301:
2296:
2291:
2286:
2280:
2278:
2271:
2270:
2268:
2267:
2262:
2261:
2260:
2255:
2245:
2240:
2235:
2229:
2227:
2221:
2220:
2218:
2217:
2216:
2215:
2210:
2200:
2188:
2187:
2186:
2185:
2180:
2175:
2170:
2165:
2160:
2155:
2150:
2145:
2140:
2135:
2130:
2125:
2120:
2115:
2110:
2100:
2095:
2090:
2085:
2080:
2075:
2066:
2064:
2058:
2057:
2055:
2054:
2053:
2052:
2047:
2037:
2032:
2027:
2021:
2019:
2013:
2012:
1997:Endopeptidases
1995:
1993:
1992:
1985:
1978:
1970:
1964:
1963:
1953:
1941:
1940:External links
1938:
1936:
1935:
1906:(4): 565–574.
1886:
1853:
1807:
1766:
1717:
1679:
1653:
1596:
1577:(3): 897–903.
1561:
1526:Nat. Cell Biol
1512:
1493:(8): 2377–85.
1476:
1441:
1405:
1363:
1322:
1301:2027.42/116965
1262:
1219:
1190:(2): 227–234.
1170:
1133:(2): 387–408.
1113:
1078:
1021:
980:
933:
882:
847:
820:J. Med. Invest
800:
751:
722:(3): 287–308.
701:
699:
696:
675:) is from the
659:
656:
631:co-polymerized
627:polyacrylamide
615:
612:
592:osteoarthritis
579:
576:
559:
556:
531:
528:
511:
508:
504:ovarian cancer
480:
477:
460:
457:
456:
455:
436:
431:
426:
421:
415:
410:
405:
400:
395:
372:
369:
367:
366:
360:
354:
348:
342:
336:
330:
324:
318:
312:
306:
300:
290:
284:
274:
263:
261:
260:Classification
258:
205:
204:
201:
200:
195:
189:
188:
175:
169:
168:
158:
151:
150:
142:
141:
128:
122:
121:
116:
110:
109:
104:
98:
97:
92:
86:
85:
80:
74:
73:
68:
61:
60:
55:
49:
48:
45:
41:
40:
36:
35:
32:
24:
23:
15:
13:
10:
9:
6:
4:
3:
2:
3250:
3239:
3236:
3234:
3231:
3229:
3226:
3224:
3221:
3220:
3218:
3208:
3203:
3198:
3194:
3181:
3180:
3179:dense granule
3176:
3171:
3170:
3167:
3157:
3154:
3152:
3149:
3148:
3146:
3144:
3140:
3134:
3131:
3129:
3126:
3124:
3121:
3119:
3116:
3114:
3111:
3110:
3108:
3106:
3102:
3096:
3093:
3091:
3088:
3086:
3085:NADPH oxidase
3083:
3081:
3078:
3076:
3073:
3071:
3068:
3067:
3065:
3063:
3059:
3056:
3053:
3049:
3043:
3040:
3038:
3035:
3033:
3030:
3027:
3023:
3019:
3017:
3014:
3012:
3009:
3008:
3006:
3003:
2999:
2995:
2991:
2983:
2978:
2976:
2971:
2969:
2964:
2963:
2960:
2948:
2945:
2943:
2940:
2938:
2935:
2933:
2930:
2928:
2925:
2923:
2920:
2918:
2915:
2913:
2910:
2909:
2907:
2903:
2897:
2894:
2892:
2889:
2887:
2884:
2882:
2879:
2877:
2873:
2870:
2868:
2865:
2863:
2860:
2858:
2855:
2853:
2850:
2848:
2845:
2844:
2842:
2840:
2836:
2830:
2827:
2825:
2822:
2820:
2817:
2815:
2812:
2810:
2807:
2805:
2802:
2800:
2797:
2795:
2792:
2790:
2787:
2785:
2782:
2780:
2777:
2775:
2772:
2770:
2767:
2765:
2762:
2760:
2757:
2756:
2754:
2752:
2748:
2742:
2739:
2737:
2734:
2732:
2729:
2727:
2724:
2723:
2721:
2719:Fruit-derived
2717:
2711:
2708:
2706:
2703:
2701:
2698:
2696:
2693:
2691:
2688:
2686:
2683:
2681:
2678:
2676:
2673:
2671:
2668:
2666:
2663:
2661:
2658:
2656:
2653:
2651:
2648:
2647:
2645:
2643:
2639:
2634:
2631:
2627:
2623:
2616:
2611:
2609:
2604:
2602:
2597:
2596:
2593:
2581:
2578:
2576:
2573:
2572:
2570:
2568:
2564:
2558:
2555:
2554:
2552:
2548:
2542:
2539:
2537:
2534:
2533:
2531:
2527:
2519:
2516:
2514:
2511:
2510:
2509:
2506:
2504:
2501:
2499:
2496:
2494:
2491:
2489:
2486:
2485:
2483:
2479:
2474:
2471:
2467:
2463:
2456:
2451:
2449:
2444:
2442:
2437:
2436:
2433:
2419:
2416:
2414:
2411:
2410:
2409:
2406:
2404:
2403:Streptokinase
2401:
2399:
2395:
2392:
2389:
2385:
2381:
2378:
2376:
2373:
2371:
2368:
2364:
2361:
2359:
2356:
2355:
2354:
2351:
2349:
2346:
2344:
2341:
2339:
2336:
2335:
2333:
2329:
2323:
2320:
2318:
2315:
2314:
2312:
2310:
2306:
2300:
2297:
2295:
2292:
2290:
2287:
2285:
2282:
2281:
2279:
2277:
2276:immune system
2272:
2266:
2265:C3-convertase
2263:
2259:
2256:
2254:
2251:
2250:
2249:
2246:
2244:
2241:
2239:
2236:
2234:
2231:
2230:
2228:
2226:
2222:
2214:
2211:
2209:
2206:
2205:
2204:
2201:
2199:
2196:
2194:
2190:
2189:
2184:
2181:
2179:
2176:
2174:
2171:
2169:
2166:
2164:
2161:
2159:
2156:
2154:
2151:
2149:
2146:
2144:
2141:
2139:
2136:
2134:
2131:
2129:
2126:
2124:
2121:
2119:
2116:
2114:
2111:
2109:
2106:
2105:
2104:
2101:
2099:
2096:
2094:
2091:
2089:
2086:
2084:
2081:
2079:
2076:
2074:
2071:
2068:
2067:
2065:
2063:
2059:
2051:
2048:
2046:
2043:
2042:
2041:
2038:
2036:
2033:
2031:
2028:
2026:
2023:
2022:
2020:
2018:
2014:
2009:
2006:
2002:
1998:
1991:
1986:
1984:
1979:
1977:
1972:
1971:
1968:
1961:
1957:
1954:
1952:
1948:
1944:
1943:
1939:
1931:
1927:
1922:
1917:
1913:
1909:
1905:
1901:
1897:
1890:
1887:
1881:
1876:
1872:
1868:
1864:
1857:
1854:
1849:
1845:
1841:
1837:
1833:
1829:
1825:
1821:
1814:
1812:
1808:
1803:
1799:
1794:
1789:
1786:(2): 853–60.
1785:
1781:
1780:J. Biol. Chem
1777:
1770:
1767:
1762:
1758:
1753:
1748:
1744:
1740:
1736:
1732:
1728:
1721:
1718:
1713:
1709:
1705:
1701:
1697:
1693:
1692:Anal. Biochem
1686:
1684:
1680:
1668:
1664:
1657:
1654:
1649:
1645:
1641:
1637:
1633:
1629:
1624:
1619:
1615:
1611:
1607:
1600:
1597:
1592:
1588:
1584:
1580:
1576:
1572:
1565:
1562:
1557:
1553:
1548:
1543:
1539:
1535:
1531:
1527:
1523:
1516:
1513:
1508:
1504:
1500:
1496:
1492:
1488:
1480:
1477:
1472:
1468:
1464:
1460:
1457:(2): 315–23.
1456:
1452:
1445:
1442:
1437:
1433:
1429:
1425:
1421:
1417:
1409:
1406:
1401:
1397:
1392:
1387:
1384:(1): 98–107.
1383:
1379:
1375:
1367:
1364:
1359:
1355:
1350:
1345:
1341:
1337:
1333:
1326:
1323:
1318:
1314:
1310:
1306:
1302:
1298:
1293:
1288:
1285:(2): 129–34.
1284:
1280:
1273:
1266:
1263:
1258:
1254:
1249:
1244:
1240:
1236:
1235:
1230:
1223:
1220:
1215:
1211:
1206:
1201:
1197:
1193:
1189:
1185:
1181:
1174:
1171:
1166:
1162:
1157:
1152:
1148:
1144:
1140:
1136:
1132:
1128:
1124:
1117:
1114:
1109:
1105:
1101:
1097:
1093:
1089:
1082:
1079:
1074:
1070:
1065:
1060:
1056:
1052:
1048:
1044:
1040:
1036:
1032:
1025:
1022:
1017:
1013:
1008:
1003:
1000:(6): 679–91.
999:
995:
991:
984:
981:
976:
972:
968:
964:
960:
956:
952:
948:
944:
937:
934:
929:
925:
920:
915:
910:
905:
901:
897:
893:
886:
883:
878:
874:
870:
866:
862:
858:
851:
848:
843:
839:
834:
829:
825:
821:
814:
807:
805:
801:
796:
792:
787:
782:
778:
774:
770:
766:
762:
755:
752:
747:
743:
739:
735:
730:
725:
721:
717:
713:
706:
703:
697:
695:
693:
689:
684:
682:
678:
674:
670:
665:
657:
655:
652:
648:
644:
640:
636:
632:
628:
624:
621:is a type of
620:
613:
611:
609:
605:
601:
597:
593:
589:
585:
577:
575:
573:
569:
565:
557:
555:
553:
549:
545:
541:
537:
529:
527:
525:
521:
517:
509:
507:
505:
501:
497:
493:
489:
485:
478:
476:
474:
470:
466:
458:
453:
449:
445:
441:
437:
435:
432:
430:
427:
425:
422:
419:
416:
414:
411:
409:
406:
404:
401:
399:
396:
393:
389:
386:
385:
384:
382:
378:
370:
364:
361:
358:
355:
352:
349:
346:
343:
340:
337:
334:
331:
328:
325:
322:
319:
316:
313:
310:
307:
304:
301:
298:
294:
291:
288:
285:
282:
278:
275:
272:
268:
265:
264:
259:
257:
255:
251:
247:
243:
239:
234:
230:
226:
222:
218:
215:
214:Ancient Greek
211:
199:
196:
194:
190:
187:
183:
179:
176:
174:
170:
166:
162:
159:
156:
152:
147:
143:
140:
136:
132:
129:
127:
123:
120:
117:
115:
111:
108:
105:
103:
99:
96:
93:
91:
87:
84:
81:
79:
75:
72:
69:
66:
62:
59:
56:
54:
50:
46:
42:
37:
30:
25:
20:
3172:
3112:
3095:Cathelicidin
3026:Proteinase 3
2994:granulocytes
2838:
2566:
2507:
2407:
2193:fibrinolysis
2191:
2069:
2035:Chymotrypsin
1903:
1899:
1889:
1870:
1866:
1856:
1823:
1819:
1783:
1779:
1769:
1734:
1730:
1720:
1695:
1691:
1672:September 1,
1670:. Retrieved
1666:
1656:
1613:
1609:
1599:
1574:
1570:
1564:
1532:(8): 970–7.
1529:
1525:
1515:
1490:
1487:Biochemistry
1486:
1479:
1454:
1450:
1444:
1419:
1415:
1408:
1381:
1377:
1366:
1339:
1335:
1325:
1282:
1278:
1265:
1238:
1232:
1222:
1187:
1184:Theranostics
1183:
1173:
1130:
1126:
1116:
1094:(3): 383–8.
1091:
1088:Exp. Eye Res
1087:
1081:
1038:
1034:
1024:
997:
993:
983:
950:
946:
936:
899:
895:
885:
860:
857:Physiol. Rev
856:
850:
826:(1–2): 1–9.
823:
819:
771:(1): 68–88.
768:
764:
754:
719:
715:
705:
685:
681:Max Bergmann
661:
639:denaturation
625:that uses a
617:
596:chronic pain
588:osteoporosis
581:
561:
552:shear stress
540:osteoporosis
533:
513:
496:amyloid beta
482:
462:
434:Pancreatitis
381:drug targets
374:
339:Cathepsin L2
333:Cathepsin L1
224:
220:
216:
209:
208:
3090:Collagenase
3075:Lactoferrin
3042:Collagenase
2947:Gingipain K
2942:Gingipain R
2912:Clostripain
2557:Nepenthesin
2098:Factor XIIa
2078:Factor VIIa
2062:Coagulation
1737:(1): 52–7.
1698:(1): 91–8.
1616:(10): 669.
1378:Circulation
1342:(1): 60–4.
558:Cathepsin V
544:Osteoclasts
536:collagenase
530:Cathepsin K
520:fibronectin
510:Cathepsin D
494:to produce
490:, cleaving
484:Cathepsin B
479:Cathepsin B
459:Cathepsin A
440:keratoconus
363:Cathepsin Z
357:Cathepsin W
351:Cathepsin S
345:Cathepsin O
327:Cathepsin K
321:Cathepsin H
315:Cathepsin G
309:Cathepsin F
303:Cathepsin E
293:Cathepsin D
287:Cathepsin C
277:Cathepsin B
267:Cathepsin A
250:osteoclasts
246:cathepsin K
219:"down" and
39:Identifiers
3238:Cathepsins
3217:Categories
3105:Eosinophil
3062:Neutrophil
2927:Autophagin
2741:Actinidain
2710:Caspase 14
2705:Caspase 13
2700:Caspase 12
2695:Caspase 10
2536:Plasmepsin
2529:Pathogenic
2481:Vertebrate
2380:Subtilisin
2322:Batroxobin
2103:Kallikrein
2093:Factor XIa
2083:Factor IXa
2050:Pancreatic
2045:Neutrophil
1956:Cathepsins
1336:Cell Cycle
698:References
619:Zymography
578:Inhibitors
566:(IR), and
242:organelles
210:Cathepsins
161:structures
3228:Proteases
3173:see also
3156:Histamine
3113:Cathepsin
3016:Defensins
2932:Cruzipain
2839:Cathepsin
2736:Bromelain
2690:Caspase 9
2685:Caspase 8
2680:Caspase 7
2675:Caspase 6
2670:Caspase 5
2665:Caspase 4
2660:Caspase 3
2655:Caspase 2
2650:Caspase 1
2622:Proteases
2567:Cathepsin
2462:Proteases
2408:Cathepsin
2394:Sedolisin
2370:Prostasin
2088:Factor Xa
1873:: 35–46.
1632:1474-1784
1279:FEBS Lett
1147:1875-8908
643:leupeptin
413:Arthritis
238:lysosomes
229:proteases
107:PDOC00126
83:IPR000668
22:Cathepsin
3143:Basophil
3080:Lysozyme
3032:Lysozyme
3020:neutral
2990:granules
2922:Separase
2493:Chymosin
2309:Venombin
2294:Tryptase
2289:Granzyme
2243:Factor I
2238:Factor D
2233:Factor B
2073:Thrombin
2070:factors:
2040:Elastase
1930:19873011
1848:17748131
1802:15393803
1761:21982919
1712:20206119
1648:10186583
1640:27681784
1591:15913550
1556:17643114
1507:10029531
1471:17305509
1436:17098827
1400:16365196
1358:17245112
1317:28099876
1257:15456852
1214:22400064
1165:21613740
1108:16893541
1073:15831716
1016:15963783
994:Immunity
975:39292302
967:23499711
953:: 1–23.
928:24282339
877:10508238
842:15751268
795:22024571
738:25436581
692:bromelin
608:COVID-19
600:STI-1558
548:collagen
452:glaucoma
178:RCSB PDB
78:InterPro
3207:Biology
3151:Heparin
2751:Calpain
2642:Caspase
2348:Pronase
2338:Acrosin
2284:Chymase
2198:Plasmin
2030:Trypsin
1951:A01.010
1921:2141516
1828:Bibcode
1820:Science
1752:3221864
1547:3065497
1309:7805878
1205:3296470
1156:4317342
1064:4797943
1043:Bibcode
1035:Science
919:3824312
786:7105208
746:2112616
669:MEDLINE
658:History
635:gelatin
524:laminin
469:lysates
392:mitogen
233:enzymes
221:hepsein
102:PROSITE
95:Pept_C1
58:PF00112
3223:EC 3.4
3193:Portal
2829:CAPNS2
2824:CAPNS1
2819:CAPN14
2814:CAPN13
2809:CAPN12
2804:CAPN11
2799:CAPN10
2731:Ficain
2726:Papain
2633:3.4.22
2488:Pepsin
2473:3.4.23
2375:Reelin
2317:Ancrod
2274:Other
2008:3.4.21
1962:(MeSH)
1947:MEROPS
1928:
1918:
1846:
1800:
1759:
1749:
1710:
1646:
1638:
1630:
1589:
1554:
1544:
1505:
1469:
1434:
1398:
1356:
1315:
1307:
1255:
1212:
1202:
1163:
1153:
1145:
1106:
1071:
1061:
1014:
973:
965:
926:
916:
875:
840:
793:
783:
744:
736:
688:papain
673:PubMed
641:using
594:, and
450:, and
398:Stroke
388:Cancer
227:) are
193:PDBsum
167:
157:
139:SUPFAM
114:MEROPS
71:CL0125
44:Symbol
2905:Other
2794:CAPN9
2789:CAPN8
2784:CAPN7
2779:CAPN6
2774:CAPN5
2769:CAPN3
2764:CAPN2
2759:CAPN1
2550:Plant
2498:Renin
2384:Furin
2331:Other
2258:MASP2
2253:MASP1
2183:KLK15
2178:KLK14
2173:KLK13
2168:KLK12
2163:KLK11
2158:KLK10
1644:S2CID
1313:S2CID
1275:(PDF)
971:S2CID
816:(PDF)
742:S2CID
418:Ebola
217:kata-
135:SCOPe
126:SCOP2
90:SMART
3054:(2°)
3004:(1°)
2398:TPP1
2248:MASP
2153:KLK9
2148:KLK8
2143:KLK7
2138:KLK6
2133:KLK5
2128:KLK4
2123:KLK3
2118:KLK2
2113:KLK1
1945:The
1926:PMID
1844:PMID
1798:PMID
1757:PMID
1708:PMID
1674:2022
1636:PMID
1628:ISSN
1587:PMID
1552:PMID
1503:PMID
1467:PMID
1432:PMID
1396:PMID
1354:PMID
1305:PMID
1253:PMID
1210:PMID
1161:PMID
1143:ISSN
1104:PMID
1069:PMID
1012:PMID
963:PMID
924:PMID
900:2013
873:PMID
838:PMID
791:PMID
769:1824
734:PMID
629:gel
522:and
424:COPD
186:PDBj
182:PDBe
165:ECOD
155:Pfam
131:1aec
67:clan
65:Pfam
53:Pfam
2992:of
2388:S1P
2108:PSA
1916:PMC
1908:doi
1875:doi
1871:119
1836:doi
1788:doi
1784:181
1747:PMC
1739:doi
1735:516
1700:doi
1696:401
1618:doi
1579:doi
1575:332
1542:PMC
1534:doi
1495:doi
1459:doi
1424:doi
1420:292
1386:doi
1382:113
1344:doi
1297:hdl
1287:doi
1283:357
1243:doi
1239:117
1200:PMC
1192:doi
1151:PMC
1135:doi
1096:doi
1059:PMC
1051:doi
1039:308
1002:doi
955:doi
951:105
914:PMC
904:doi
865:doi
828:doi
781:PMC
773:doi
724:doi
604:FDA
252:in
225:CTS
173:PDB
47:CTP
3219::
3177:,
2876:L2
2872:L1
2630:EC
2624::
2470:EC
2464::
2005:EC
1999::
1924:.
1914:.
1904:20
1902:.
1898:.
1869:.
1865:.
1842:.
1834:.
1824:84
1822:.
1810:^
1796:.
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119:C1
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2896:Z
2891:W
2886:S
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2867:K
2862:H
2857:F
2852:C
2847:B
2635:)
2628:(
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2607:t
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2580:E
2575:D
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1989:e
1982:t
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