600:
668:
2316:
648:
575:
380:
C6-C9 all contain a MACPF domain and assemble into the membrane attack complex. C6, C7 and C8β appear to be non-lytic and function as scaffold proteins within the MAC. In contrast both C8α and C9 are capable of lysing cells. The final stage of MAC formation involves polymerisation of C9 into a large
391:
is stored in granules within cytotoxic T-cells and is responsible for killing virally infected and transformed cells. Perforin functions via two distinct mechanisms. Firstly, like C9, high concentrations of perforin can form pores that lyse cells. Secondly, perforin permits delivery of the cytotoxic
627:
inhibitors and prevent inappropriate activity of complement against self cells (Figure 3). Biochemical studies have revealed the peptide sequences in C8α and C9 that bind to CD59. Analysis of the MACPF domain structures reveals that these sequences map to the second cluster of helices that unfurl
339:
Proteins containing MACPF domains play key roles in vertebrate immunity, embryonic development, and neural-cell migration. The ninth component of complement and perforin form oligomeric pores that lyse bacteria and kill virus-infected cells, respectively. The crystal structure of a bacterial MACPF
1455:
Tsutsui, Tomokazu; Morita-Yamamuro, Chizuko; Asada, Yutaka; Minami, Eiichi; Shibuya, Naoto; Ikeda, Akira; Yamaguchi, Junji (1 September 2006). "Salicylic acid and a chitin elicitor both control expression of the CAD1 gene involved in the plant immunity of
Arabidopsis".
513:
It is suggested that MACPF proteins and CDCs form pores in the same way (figure 1). Specifically it is hypothesised that MACPF proteins oligomerise to form a large circular pore (figure 2). A concerted conformational change within each monomer then results in two
2106:
Kieffer B, Driscoll PC, Campbell ID, Willis AC, van der Merwe PA, Davis SJ (1994). "Three-dimensional solution structure of the extracellular region of the complement regulatory protein CD59, a new cell-surface protein domain related to snake venom neurotoxins".
2070:
Lovelace LL, Chiswell B, Slade DJ, Sodetz JM, Lebioda L (2007). "Crystal structure of complement protein C8gamma in complex with a peptide containing the C8gamma binding site on C8alpha: Implications for C8gamma ligand binding".
451:), which controls embryonic patterning, also contains a MACPF domain. Its function is implicated in a receptor tyrosine kinase signaling pathway that specifies differentiation and terminal cell fate.
1603:
Ishino T, Chinzei Y, Yuda M (2005). "A Plasmodium sporozoite protein with a membrane attack complex domain is required for breaching the liver sinusoidal cell layer prior to hepatocyte infection".
1517:
Oshiro N, Kobayashi C, Iwanaga S, et al. (2004). "A new membrane-attack complex/perforin (MACPF) domain lethal toxin from the nematocyst venom of the
Okinawan sea anemone Actineria villosa".
891:
Rosado, Carlos J.; Kondos, Stephanie; Bull, Tara E.; Kuiper, Michael J.; Law, Ruby H. P.; Buckle, Ashley M.; Voskoboinik, Ilia; Bird, Phillip I.; Trapani, Joseph A. (1 September 2008).
1300:
Rosado, Carlos J.; Buckle, Ashley M.; Law, Ruby H. P.; Butcher, Rebecca E.; Kan, Wan-Ting; Bird, Catherina H.; Ung, Kheng; Browne, Kylie A.; Baran, Katherine (14 September 2007).
362:, suggesting that MACPF proteins create pores and disrupt cell membranes similar to cytolysin. A representative list of proteins belonging to the MACPF family can be found in the
1089:
Witzel-Schlömp K, Späth PJ, Hobart MJ, et al. (1997). "The human complement C9 gene: identification of two mutations causing deficiency and revision of the gene structure".
766:
628:
to span the membrane. It is therefore suggested that CD59 directly inhibits the MAC by interfering with conformational change in one of the membrane spanning regions.
2142:
Kira R, Ihara K, Takada H, Gondo K, Hara T (1998). "Nonsense mutation in exon 4 of human complement C9 gene is the major cause of
Japanese complement C9 deficiency".
187:
948:
Tschopp J, Masson D, Stanley KK (1986). "Structural/functional similarity between proteins involved in complement- and cytotoxic T-lymphocyte-mediated cytolysis".
510:
sequence identity between the two families is extremely low, and the relationship is not detectable using conventional sequence based data mining techniques.
1204:
Michael A. Hadders; Dennis X. Beringer & Piet Gros (2007). "Structure of C8-MACPF Reveals
Mechanism of Membrane Attack in Complement Immune Defense".
701:. Overactivity of MACPF proteins can also cause disease. Most notably, deficiency of the MAC inhibitor CD59 results in an overactivity of complement and
2020:"Expression and characterization of recombinant subunits of human complement component C8: further analysis of the function of C8 alpha and C8 gamma"
1050:"Perforin activity and immune homeostasis: the common A91V polymorphism in perforin results in both presynaptic and postsynaptic defects in function"
599:
131:
119:
635:
family and interacts with C8α. The binding site on C8α is known, however, the precise role of C8γ in the MAC remains to be understood.
2223:
Verbsky JW, Grossman WJ (2006). "Hemophagocytic lymphohistiocytosis: diagnosis, pathophysiology, treatment, and future perspectives".
1416:"The Arabidopsis gene CAD1 controls programmed cell death in the plant immune system and encodes a protein containing a MACPF domain"
83:
712:
762:
758:
667:
750:
746:
1646:
Zheng C, Heintz N, Hatten ME (1996). "CNS gene encoding astrotactin, which supports neuronal migration along glial fibers".
207:
2363:
2353:
1738:
Martin JR, Raibaud A, Ollo R (1994). "Terminal pattern elements in
Drosophila embryo induced by the torso-like protein".
1257:
Anderluh, Gregor; Lakey, Jeremy H. (1 October 2008). "Disparate proteins use similar architectures to damage membranes".
702:
392:
granzymes A and B into target cells. Once delivered, granzymes are able to induce apoptosis and cause target cell death.
289:. Deficiency of either protein can result in human disease. Structural studies reveal that MACPF domains are related to
231:
20:
529:
of the target cell. Like CDC's MACPF proteins are thus β-pore forming toxins that act like a molecular hole punch.
788:
Gilbert, Robert J. C.; Mikelj, Miha; Dalla Serra, Mauro; Froelich, Christopher J.; Anderluh, Gregor (1 June 2013).
2348:
1699:"Apextrin, a novel extracellular protein associated with larval ectoderm evolution in Heliocidaris erythrogramma"
355:
313:
290:
690:
Deficiency of C9, or other components of the MAC results in an increased susceptibility to diseases caused by
195:
281:
virally infected and transformed cells. In addition, perforin permits delivery of cytotoxic proteases called
1002:
Voskoboinik I, Smyth MJ, Trapani JA (2006). "Perforin-mediated target-cell death and immune homeostasis".
2358:
454:
Functionally uncharacterised MACPF proteins are sporadically distributed in bacteria. Several species of
2343:
2306:
437:
191:
88:
2338:
481:
460:
2266:
Wright KO, Messing EM, Reeder JE (2004). "DBCCR1 mediates death in cultured bladder tumor cells".
1979:"Identity of the segment of human complement C8 recognized by complement regulatory protein CD59"
624:
498:
1840:"Structure of a cholesterol-binding, thiol-activated cytolysin and a model of its membrane form"
606:
Molecular model of the pre-pore form of a MACPF protein based upon the structure of pneunolysin.
754:
742:
738:
382:
359:
258:
239:
675:
659:
586:
583:
486:
374:
Many proteins belonging to the MACPF superfamily play key roles in plant and animal immunity.
853:
Peitsch MC, Tschopp J (1991). "Assembly of macromolecular pores by immune defense systems".
1747:
1655:
1313:
1160:
957:
174:
532:
Other crystal structures for members of the MACPF superfamily can be found in RCSB: i.e.,
8:
489:
294:
251:
223:
143:
1751:
1659:
1317:
1164:
961:
822:
789:
2291:
2248:
2167:
2049:
1918:
1869:
1820:
1771:
1679:
1628:
1499:
1391:
1366:
1347:
1239:
1186:
1114:
1027:
981:
925:
892:
835:
448:
410:
396:
266:
124:
1856:
1839:
1807:
1790:
1580:
1553:
432:
428:
2283:
2240:
2205:
2159:
2124:
2088:
2041:
2000:
1959:
1910:
1861:
1812:
1763:
1720:
1683:
1671:
1620:
1616:
1585:
1554:"Essential role of membrane-attack protein in malarial transmission to mosquito host"
1534:
1491:
1483:
1437:
1396:
1339:
1331:
1282:
1274:
1231:
1178:
1106:
1071:
1019:
973:
930:
912:
908:
870:
866:
827:
809:
377:
346:
274:
235:
182:
2252:
2053:
1632:
1503:
1351:
1243:
1190:
1118:
1031:
839:
363:
51:
2295:
2275:
2232:
2197:
2171:
2151:
2116:
2080:
2031:
1990:
1949:
1922:
1900:
1873:
1851:
1824:
1802:
1775:
1755:
1710:
1663:
1612:
1575:
1565:
1526:
1473:
1465:
1427:
1386:
1378:
1321:
1266:
1221:
1213:
1168:
1098:
1061:
1011:
985:
965:
920:
904:
862:
817:
801:
170:
1530:
324:
318:
309:
148:
136:
112:
2084:
1667:
1367:"The major human and mouse granzymes are structurally and functionally divergent"
1102:
1066:
1049:
997:
995:
76:
2201:
2320:
2036:
2019:
1905:
1888:
1270:
727:
477:
305:
As of early 2016, there are three families belonging to the MACPF superfamily:
247:
227:
64:
2236:
992:
805:
2332:
1487:
1335:
1278:
1203:
916:
813:
695:
691:
631:
Other proteins that bind to the MAC include C8γ. This protein belongs to the
526:
493:
328:
270:
1570:
1326:
1301:
1217:
1173:
1148:
2287:
2279:
2244:
2209:
2185:
2092:
1995:
1978:
1963:
1954:
1937:
1914:
1816:
1724:
1715:
1698:
1624:
1589:
1538:
1495:
1441:
1400:
1343:
1286:
1235:
1182:
1075:
1023:
934:
831:
503:
2163:
2155:
2128:
2045:
2004:
1865:
1767:
1675:
1382:
1110:
977:
874:
1432:
1415:
716:
589:
519:
515:
464:
also contains a MACPF protein, however, this molecule appears non-lytic.
424:
403:
254:. In eukaryotes, MACPF proteins play a role in immunity and development.
2120:
1977:
Lockert DH, Kaufman KM, Chang CP, Hüsler T, Sodetz JM, Sims PJ (1995).
1889:"Structural basis of pore formation by the bacterial toxin pneumolysin"
698:
522:
507:
443:
369:
286:
226:, sometimes referred to as the MACPF/CDC superfamily, is named after a
100:
1478:
1469:
1226:
558:
552:
546:
540:
534:
350:
1759:
969:
715:(FHL or HLH). This disease is characterised by an overactivation of
647:
632:
1015:
574:
423:
Not all MACPF proteins function in defence or attack. For example,
354:). The MACPF domain is structurally similar to pore-forming
2315:
720:
708:
417:
388:
282:
262:
243:
71:
334:
1454:
399:) functions in the plant immune response to bacterial infection.
95:
1838:
Rossjohn J, Feil SC, McKinstry WJ, Tweten RK, Parker MW (1997).
1302:"A common fold mediates vertebrate defense and bacterial attack"
1149:"A Common Fold Mediates Vertebrate Defense and Bacterial Attack"
787:
592:
that are proposed to unwind and pierce the membrane are in pink.
431:) is involved in neural cell migration in mammals and apextrin (
314:
The Thiol-activated
Cholesterol-dependent Cytolysin (CDC) Family
1887:
Tilley SJ, Orlova EV, Gilbert RJ, Andrew PW, Saibil HR (2005).
726:
The MACPF protein DBCCR1 may function as a tumor suppressor in
674:
Crystal structure of C8γ (green) with peptide from C8α (cyan).
473:
202:
2105:
472:
The X-ray crystal structure of Plu-MACPF, a protein from the
278:
1551:
2069:
1837:
1552:
Kadota K, Ishino T, Matsuyama T, Chinzei Y, Yuda M (2004).
770:
655:
620:
164:
107:
58:
46:
2018:
Schreck SF, Plumb ME, Platteborze PL, et al. (1998).
1936:
Huang Y, Qiao F, Abagyan R, Hazard S, Tomlinson S (2006).
1791:"Chlamydial homologues of the MACPF (MAC/perforin) domain"
1414:
Morita-Yamamuro C, Tsutsui T, Sato M, et al. (2005).
1413:
416:
MACPF proteins are also important for the invasion of the
1976:
1886:
1088:
1048:
Voskoboinik I, Sutton VR, Ciccone A, et al. (2007).
488:
These data reveal that the MACPF domain is homologous to
1047:
733:
458:
contain MACPF proteins. The insect pathogenic bacteria
2017:
1001:
1935:
1516:
1299:
890:
269:. C9 functions by punching holes in the membranes of
2304:
329:
321:- The Membrane Attack Complex/Perforin (MACPF) Family
2141:
947:
370:
Biological roles of MACPF domain containing proteins
2265:
1696:
1147:Rosado CJ, Buckle AM, Law RH, et al. (2007).
790:"Effects of MACPF/CDC proteins on lipid membranes"
711:deficiency results in the commonly fatal disorder
381:pore that punches a hole in the outer membrane of
1737:
1645:
1602:
1365:Kaiserman D, Bird CH, Sun J, et al. (2006).
2330:
1364:
2222:
1146:
582:: a) The structure of the CDC perfringolysin O
420:parasite into the mosquito host and the liver.
335:Membrane Attack Complex/Perforin (MACPF) Family
852:
588:. In both proteins the two small clusters of
567:Mechanism of membrane attack by MACPF proteins
492:cholesterol dependent cytolysins (CDC's) from
297:previously thought to only exist in bacteria.
1697:Haag ES, Sly BJ, Andrews ME, Raff RA (1999).
1256:
893:"The MACPF/CDC family of pore-forming toxins"
614:
1458:Bioscience, Biotechnology, and Biochemistry
713:familial hemophagocytic lymphohistiocytosis
2188:(2001). "Complement. First of two parts".
2065:
2063:
1938:"Defining the CD59-C9 binding interaction"
1788:
1142:
1140:
1138:
1136:
1134:
1132:
1130:
1128:
467:
2035:
1994:
1953:
1904:
1855:
1806:
1714:
1579:
1569:
1477:
1431:
1390:
1325:
1225:
1172:
1065:
924:
821:
685:
666:
646:
598:
573:
221:Membrane Attack Complex/Perforin (MACPF)
2184:
2060:
1125:
619:Complement regulatory proteins such as
2331:
1043:
1041:
734:Human proteins containing this domain
257:Archetypal members of the family are
886:
884:
794:Cellular and Molecular Life Sciences
703:Paroxysmal nocturnal hemoglobinuria
364:Transporter Classification Database
13:
1038:
585:and b) the structure of Plu-MACPF
273:bacteria. Perforin is released by
265:, both of which function in human
21:Complement membrane attack complex
14:
2375:
881:
16:Large group of transport proteins
2314:
1617:10.1111/j.1462-5822.2004.00447.x
909:10.1111/j.1462-5822.2008.01191.x
640:Proteins that bind MACPF domains
356:cholesterol-dependent cytolysins
291:cholesterol-dependent cytolysins
2259:
2216:
2178:
2135:
2099:
2011:
1970:
1929:
1880:
1831:
1782:
1731:
1690:
1639:
1596:
1545:
1510:
1448:
1407:
1358:
1293:
1250:
518:regions unwinding to form four
485:has been determined (figure 1).
1259:Trends in Biochemical Sciences
1197:
1082:
941:
846:
781:
1:
1857:10.1016/S0092-8674(00)80251-2
1808:10.1016/S0960-9822(00)80102-5
1531:10.1016/j.toxicon.2003.11.017
775:
435:) is involved in sea urchin (
159:Available protein structures:
2085:10.1016/j.molimm.2007.06.359
1668:10.1126/science.272.5260.417
1558:Proc. Natl. Acad. Sci. U.S.A
1103:10.4049/jimmunol.158.10.5043
1067:10.1182/blood-2007-02-072850
867:10.1016/0955-0674(91)90045-Z
496:pathogenic bacteria such as
413:)protein as a lethal toxin.
7:
2202:10.1056/NEJM200104053441406
300:
10:
2380:
2364:Integral membrane proteins
2354:Transmembrane transporters
2037:10.4049/jimmunol.161.1.311
1906:10.1016/j.cell.2005.02.033
1271:10.1016/j.tibs.2008.07.004
665:
645:
597:
572:
438:Heliocidaris erythrogramma
252:pore-forming toxins (PFTs)
18:
2237:10.1080/07853890500465189
806:10.1007/s00018-012-1153-8
615:Control of MACPF proteins
201:
181:
163:
158:
154:
142:
130:
118:
106:
94:
82:
70:
57:
45:
37:
32:
27:
482:Photorhabdus luminescens
461:Photorhabdus luminescens
409:uses a MACPF (AvTX-60A;
395:The plant protein CAD1 (
342:Photorhabdus luminescens
340:protein, Plu-MACPF from
1571:10.1073/pnas.0406187101
1327:10.1126/science.1144706
1218:10.1126/science.1147103
1174:10.1126/science.1144706
723:mediated organ damage.
499:Clostridium perfringens
468:Structure and mechanism
232:membrane attack complex
2349:Transmembrane proteins
2280:10.1038/sj.onc.1206642
1996:10.1074/jbc.270.8.3483
1955:10.1074/jbc.M603690200
1716:10.1006/dbio.1999.9283
678:
662:
607:
594:
383:gram-negative bacteria
360:gram-positive bacteria
246:(PF). Members of this
238:(C6, C7, C8α, C8β and
230:that is common to the
2156:10.1007/s004390050749
1383:10.1083/jcb.200606073
897:Cellular Microbiology
855:Curr. Opin. Cell Biol
686:Role in human disease
670:
650:
602:
577:
236:the complement system
2073:Molecular Immunology
447:Torso-like protein (
293:(CDCs), a family of
2121:10.1021/bi00181a006
1789:Ponting CP (1999).
1752:1994Natur.367..741M
1660:1996Sci...272..417Z
1318:2007Sci...317.1548R
1312:(5844): 1548–1551.
1212:(5844): 1552–1554.
1165:2007Sci...317.1548R
962:1986Natur.322..831T
642:
569:
378:Complement proteins
295:pore forming toxins
28:MAC/Perforin domain
2359:Transport proteins
1433:10.1093/pcp/pci095
1420:Plant Cell Physiol
679:
663:
638:
608:
595:
565:
234:(MAC) proteins of
2344:Membrane proteins
1948:(37): 27398–404.
1470:10.1271/bbb.50700
1159:(5844): 1548–51.
1004:Nat. Rev. Immunol
800:(12): 2083–2098.
719:which results in
694:bacteria such as
683:
682:
654:NMR structure of
612:
611:
407:Actineria villosa
275:cytotoxic T cells
217:
216:
213:
212:
208:structure summary
2371:
2339:Protein families
2319:
2318:
2310:
2300:
2299:
2263:
2257:
2256:
2220:
2214:
2213:
2186:Walport, Mark J.
2182:
2176:
2175:
2139:
2133:
2132:
2103:
2097:
2096:
2067:
2058:
2057:
2039:
2015:
2009:
2008:
1998:
1974:
1968:
1967:
1957:
1933:
1927:
1926:
1908:
1884:
1878:
1877:
1859:
1835:
1829:
1828:
1810:
1786:
1780:
1779:
1760:10.1038/367741a0
1735:
1729:
1728:
1718:
1694:
1688:
1687:
1643:
1637:
1636:
1600:
1594:
1593:
1583:
1573:
1549:
1543:
1542:
1514:
1508:
1507:
1481:
1464:(9): 2042–2048.
1452:
1446:
1445:
1435:
1411:
1405:
1404:
1394:
1362:
1356:
1355:
1329:
1297:
1291:
1290:
1254:
1248:
1247:
1229:
1201:
1195:
1194:
1176:
1144:
1123:
1122:
1086:
1080:
1079:
1069:
1045:
1036:
1035:
999:
990:
989:
970:10.1038/322831a0
945:
939:
938:
928:
903:(9): 1765–1774.
888:
879:
878:
850:
844:
843:
825:
785:
643:
637:
570:
564:
561:
555:
549:
543:
537:
353:
344:was determined (
156:
155:
25:
24:
2379:
2378:
2374:
2373:
2372:
2370:
2369:
2368:
2329:
2328:
2325:
2313:
2305:
2303:
2264:
2260:
2221:
2217:
2196:(14): 1058–66.
2190:N. Engl. J. Med
2183:
2179:
2140:
2136:
2115:(15): 4471–82.
2104:
2100:
2068:
2061:
2016:
2012:
1989:(34): 19723–8.
1975:
1971:
1934:
1930:
1885:
1881:
1836:
1832:
1787:
1783:
1746:(6465): 741–5.
1736:
1732:
1695:
1691:
1654:(5260): 417–9.
1644:
1640:
1605:Cell. Microbiol
1601:
1597:
1564:(46): 16310–5.
1550:
1546:
1515:
1511:
1453:
1449:
1412:
1408:
1363:
1359:
1298:
1294:
1265:(10): 482–490.
1255:
1251:
1202:
1198:
1145:
1126:
1087:
1083:
1046:
1039:
1016:10.1038/nri1983
1000:
993:
956:(6082): 831–4.
946:
942:
889:
882:
851:
847:
786:
782:
778:
736:
688:
617:
557:
551:
545:
539:
533:
470:
449:TC# 1.C.39.15.1
441:) development.
411:TC# 1.C.39.10.1
397:TC# 1.C.39.11.3
372:
345:
337:
303:
120:OPM superfamily
23:
17:
12:
11:
5:
2377:
2367:
2366:
2361:
2356:
2351:
2346:
2341:
2324:
2323:
2302:
2301:
2258:
2215:
2177:
2134:
2098:
2059:
2010:
1969:
1928:
1879:
1830:
1801:(24): R911–3.
1781:
1730:
1689:
1638:
1611:(2): 199–208.
1595:
1544:
1509:
1447:
1406:
1357:
1292:
1249:
1196:
1124:
1097:(10): 5043–9.
1081:
1060:(4): 1184–90.
1037:
1010:(12): 940–52.
991:
940:
880:
845:
779:
777:
774:
735:
732:
728:bladder cancer
687:
684:
681:
680:
664:
616:
613:
610:
609:
596:
525:that span the
502:(which causes
478:enterobacteria
469:
466:
433:TC# 1.C.39.7.4
429:TC# 9.B.87.3.1
371:
368:
336:
333:
332:
331:
322:
316:
302:
299:
248:protein family
215:
214:
211:
210:
205:
199:
198:
185:
179:
178:
168:
161:
160:
152:
151:
146:
140:
139:
134:
128:
127:
122:
116:
115:
110:
104:
103:
98:
92:
91:
86:
80:
79:
74:
68:
67:
62:
55:
54:
49:
43:
42:
39:
35:
34:
30:
29:
15:
9:
6:
4:
3:
2:
2376:
2365:
2362:
2360:
2357:
2355:
2352:
2350:
2347:
2345:
2342:
2340:
2337:
2336:
2334:
2327:
2322:
2317:
2312:
2311:
2308:
2297:
2293:
2289:
2285:
2281:
2277:
2273:
2269:
2262:
2254:
2250:
2246:
2242:
2238:
2234:
2230:
2226:
2219:
2211:
2207:
2203:
2199:
2195:
2191:
2187:
2181:
2173:
2169:
2165:
2161:
2157:
2153:
2150:(6): 605–10.
2149:
2145:
2138:
2130:
2126:
2122:
2118:
2114:
2110:
2102:
2094:
2090:
2086:
2082:
2078:
2074:
2066:
2064:
2055:
2051:
2047:
2043:
2038:
2033:
2029:
2025:
2021:
2014:
2006:
2002:
1997:
1992:
1988:
1984:
1983:J. Biol. Chem
1980:
1973:
1965:
1961:
1956:
1951:
1947:
1943:
1942:J. Biol. Chem
1939:
1932:
1924:
1920:
1916:
1912:
1907:
1902:
1899:(2): 247–56.
1898:
1894:
1890:
1883:
1875:
1871:
1867:
1863:
1858:
1853:
1850:(5): 685–92.
1849:
1845:
1841:
1834:
1826:
1822:
1818:
1814:
1809:
1804:
1800:
1796:
1792:
1785:
1777:
1773:
1769:
1765:
1761:
1757:
1753:
1749:
1745:
1741:
1734:
1726:
1722:
1717:
1712:
1708:
1704:
1700:
1693:
1685:
1681:
1677:
1673:
1669:
1665:
1661:
1657:
1653:
1649:
1642:
1634:
1630:
1626:
1622:
1618:
1614:
1610:
1606:
1599:
1591:
1587:
1582:
1577:
1572:
1567:
1563:
1559:
1555:
1548:
1540:
1536:
1532:
1528:
1524:
1520:
1513:
1505:
1501:
1497:
1493:
1489:
1485:
1480:
1475:
1471:
1467:
1463:
1459:
1451:
1443:
1439:
1434:
1429:
1426:(6): 902–12.
1425:
1421:
1417:
1410:
1402:
1398:
1393:
1388:
1384:
1380:
1377:(4): 619–30.
1376:
1372:
1368:
1361:
1353:
1349:
1345:
1341:
1337:
1333:
1328:
1323:
1319:
1315:
1311:
1307:
1303:
1296:
1288:
1284:
1280:
1276:
1272:
1268:
1264:
1260:
1253:
1245:
1241:
1237:
1233:
1228:
1223:
1219:
1215:
1211:
1207:
1200:
1192:
1188:
1184:
1180:
1175:
1170:
1166:
1162:
1158:
1154:
1150:
1143:
1141:
1139:
1137:
1135:
1133:
1131:
1129:
1120:
1116:
1112:
1108:
1104:
1100:
1096:
1092:
1085:
1077:
1073:
1068:
1063:
1059:
1055:
1051:
1044:
1042:
1033:
1029:
1025:
1021:
1017:
1013:
1009:
1005:
998:
996:
987:
983:
979:
975:
971:
967:
963:
959:
955:
951:
944:
936:
932:
927:
922:
918:
914:
910:
906:
902:
898:
894:
887:
885:
876:
872:
868:
864:
860:
856:
849:
841:
837:
833:
829:
824:
819:
815:
811:
807:
803:
799:
795:
791:
784:
780:
773:
772:
768:
764:
760:
756:
752:
748:
744:
740:
731:
729:
724:
722:
718:
714:
710:
706:
704:
700:
697:
696:meningococcal
693:
692:gram-negative
676:
673:
669:
660:
657:
653:
649:
644:
641:
636:
634:
629:
626:
622:
605:
601:
593:
591:
587:
584:
581:
576:
571:
568:
563:
560:
554:
548:
542:
536:
530:
528:
524:
521:
517:
511:
509:
505:
501:
500:
495:
494:gram-positive
491:
487:
484:
483:
479:
475:
465:
463:
462:
457:
452:
450:
446:
445:
440:
439:
434:
430:
426:
425:astrotactin-1
421:
419:
414:
412:
408:
405:
400:
398:
393:
390:
386:
384:
379:
375:
367:
365:
361:
357:
352:
348:
343:
330:
326:
323:
320:
317:
315:
311:
308:
307:
306:
298:
296:
292:
288:
284:
280:
276:
272:
271:Gram-negative
268:
264:
260:
259:complement C9
255:
253:
249:
245:
241:
237:
233:
229:
225:
222:
209:
206:
204:
200:
197:
193:
189:
186:
184:
180:
176:
172:
169:
166:
162:
157:
153:
150:
147:
145:
141:
138:
135:
133:
129:
126:
123:
121:
117:
114:
111:
109:
105:
102:
99:
97:
93:
90:
87:
85:
81:
78:
75:
73:
69:
66:
63:
60:
56:
53:
50:
48:
44:
40:
36:
31:
26:
22:
2326:
2274:(1): 82–90.
2271:
2267:
2261:
2231:(1): 20–31.
2228:
2224:
2218:
2193:
2189:
2180:
2147:
2143:
2137:
2112:
2109:Biochemistry
2108:
2101:
2079:(3): 750–6.
2076:
2072:
2030:(1): 311–8.
2027:
2023:
2013:
1986:
1982:
1972:
1945:
1941:
1931:
1896:
1892:
1882:
1847:
1843:
1833:
1798:
1794:
1784:
1743:
1739:
1733:
1709:(1): 77–87.
1706:
1702:
1692:
1651:
1647:
1641:
1608:
1604:
1598:
1561:
1557:
1547:
1525:(2): 225–8.
1522:
1518:
1512:
1461:
1457:
1450:
1423:
1419:
1409:
1374:
1371:J. Cell Biol
1370:
1360:
1309:
1305:
1295:
1262:
1258:
1252:
1209:
1205:
1199:
1156:
1152:
1094:
1090:
1084:
1057:
1053:
1007:
1003:
953:
949:
943:
900:
896:
861:(4): 710–6.
858:
854:
848:
797:
793:
783:
737:
725:
707:
689:
671:
651:
639:
630:
623:function as
618:
603:
579:
578:
566:
531:
512:
504:gas gangrene
497:
490:pore forming
480:
471:
459:
455:
453:
442:
436:
422:
415:
406:
401:
394:
387:
376:
373:
341:
338:
304:
256:
220:
218:
717:lymphocytes
520:amphipathic
476:pathogenic
404:sea anemone
285:that cause
224:superfamily
132:OPM protein
33:Identifiers
2333:Categories
2144:Hum. Genet
2024:J. Immunol
1795:Curr. Biol
1479:2115/14840
1227:1874/26831
1091:J. Immunol
776:References
699:meningitis
590:α-helicesl
508:amino acid
444:Drosophila
287:cell death
171:structures
144:Membranome
19:See also:
1703:Dev. Biol
1684:206576029
1488:0916-8451
1336:1095-9203
1279:0968-0004
917:1462-5822
814:1420-9071
757:;
745:;
741:;
672:Figure 3:
652:Figure 3:
633:lipocalin
604:Figure 2:
556:,
550:,
544:,
538:,
523:β-strands
516:α-helical
456:Chlamydia
283:granzymes
101:PDOC00251
77:IPR001862
2288:14712213
2268:Oncogene
2253:30781596
2245:16448985
2225:Ann. Med
2210:11287977
2093:17692377
2054:86614986
1964:16844690
1915:15851031
1817:10608922
1725:10373306
1633:19341572
1625:15659064
1590:15520375
1539:15019483
1504:37799485
1496:16960394
1442:15799997
1401:17116752
1352:20372720
1344:17717151
1287:18778941
1244:44959101
1236:17872444
1191:20372720
1183:17717151
1119:22684521
1076:17475905
1032:39504110
1024:17124515
935:18564372
840:17084919
832:22983385
823:11114033
753:;
749:;
721:cytokine
709:Perforin
580:Figure 1
562:
527:membrane
506:). The
418:Malarial
389:Perforin
301:Families
267:immunity
263:perforin
244:perforin
188:RCSB PDB
72:InterPro
2321:Biology
2296:3620390
2172:3227108
2164:9703418
2129:7512825
2046:9647238
2005:7544344
1923:1613454
1874:8345038
1866:9182756
1825:7748405
1776:4274979
1768:8107870
1748:Bibcode
1676:8602532
1656:Bibcode
1648:Science
1519:Toxicon
1392:2064598
1314:Bibcode
1306:Science
1206:Science
1161:Bibcode
1153:Science
1111:9144525
986:4330219
978:2427956
958:Bibcode
926:2654483
875:1722985
96:PROSITE
52:PF01823
2307:Portal
2294:
2286:
2251:
2243:
2208:
2170:
2162:
2127:
2091:
2052:
2044:
2003:
1962:
1921:
1913:
1872:
1864:
1823:
1815:
1774:
1766:
1740:Nature
1723:
1682:
1674:
1631:
1623:
1588:
1581:524694
1578:
1537:
1502:
1494:
1486:
1440:
1399:
1389:
1350:
1342:
1334:
1285:
1277:
1242:
1234:
1189:
1181:
1117:
1109:
1074:
1030:
1022:
984:
976:
950:Nature
933:
923:
915:
873:
838:
830:
820:
812:
765:;
761:;
474:insect
325:1.C.97
319:1.C.39
310:1.C.12
242:) and
228:domain
203:PDBsum
177:
167:
113:1.C.39
38:Symbol
2292:S2CID
2249:S2CID
2168:S2CID
2050:S2CID
1919:S2CID
1870:S2CID
1821:S2CID
1772:S2CID
1680:S2CID
1629:S2CID
1500:S2CID
1348:S2CID
1240:S2CID
1187:S2CID
1115:S2CID
1054:Blood
1028:S2CID
982:S2CID
836:S2CID
767:MPEG1
763:FAM5C
759:FAM5B
358:from
279:lyses
89:MACPF
84:SMART
41:MACPF
2284:PMID
2241:PMID
2206:PMID
2160:PMID
2125:PMID
2089:PMID
2042:PMID
2001:PMID
1960:PMID
1911:PMID
1893:Cell
1862:PMID
1844:Cell
1813:PMID
1764:PMID
1721:PMID
1672:PMID
1621:PMID
1586:PMID
1535:PMID
1492:PMID
1484:ISSN
1438:PMID
1397:PMID
1340:PMID
1332:ISSN
1283:PMID
1275:ISSN
1232:PMID
1179:PMID
1107:PMID
1072:PMID
1020:PMID
974:PMID
931:PMID
913:ISSN
871:PMID
828:PMID
810:ISSN
771:PRF1
656:CD59
621:CD59
559:3OJY
553:3RD7
547:3QQH
541:3QOS
535:3KK7
402:The
351:2QP2
277:and
261:and
250:are
219:The
196:PDBj
192:PDBe
175:ECOD
165:Pfam
137:6h04
108:TCDB
61:clan
59:Pfam
47:Pfam
2276:doi
2233:doi
2198:doi
2194:344
2152:doi
2148:102
2117:doi
2081:doi
2032:doi
2028:161
1991:doi
1987:270
1950:doi
1946:281
1901:doi
1897:121
1852:doi
1803:doi
1756:doi
1744:367
1711:doi
1707:211
1664:doi
1652:272
1613:doi
1576:PMC
1566:doi
1562:101
1527:doi
1474:hdl
1466:doi
1428:doi
1387:PMC
1379:doi
1375:175
1322:doi
1310:317
1267:doi
1222:hdl
1214:doi
1210:317
1169:doi
1157:317
1099:doi
1095:158
1062:doi
1058:110
1012:doi
966:doi
954:322
921:PMC
905:doi
863:doi
818:PMC
802:doi
751:C8B
747:C8A
625:MAC
347:PDB
183:PDB
149:233
125:168
65:CDC
2335::
2290:.
2282:.
2272:23
2270:.
2247:.
2239:.
2229:38
2227:.
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2166:.
2158:.
2146:.
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2026:.
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1999:.
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1981:.
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1940:.
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1462:70
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739:C6
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385:.
366:.
349::
327:-
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240:C9
194:;
190:;
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2174:.
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427:(
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