20:
1868:
513:
through the CARD domain. All these protein-protein interaction form a complex called the inflammasome. The aggregation of the pro-caspase-1 causes the autocleavage and formation of an active enzyme. Caspase-1 is important for the proteolytic processing of the pro-inflammatory cytokines
122:(LRR) and variable N-terminal interaction domain. NACHT domain mediates ATP-dependent self-oligomerization and LRR senses the presence of ligand. N-terminal domain is responsible for homotypic protein-protein interaction and it can consist of
1305:
Okafuji I, Nishikomori R, Kanazawa N, Kambe N, Fujisawa A, Yamazaki S, et al. (2009). "Role of the NOD2 genotype in the clinical phenotype of Blau syndrome and early-onset sarcoidosis".
393:
The well-described receptors are NOD1 and NOD2. The recognition of their ligands recruits oligomerization of NACHT domain and CARD-CARD interaction with CARD-containing serine-threonin
1516:
146:
in 2008. The family was characterized as NLRs to provide description of the families features – NLR means nucleotide-binding domain and leucine-rich repeat containing gene family.
481:
NLRPs subfamily contains NLRP1-NLRP14 that are characterized by the presence of PYD domain. IPAF subfamily has two members – IPAF with CARD domain and NAIP with BIR domain.
1677:
1509:
385:
NODs subfamily consists of NOD1, NOD2, NOD3, NOD4 with CARD domain, CIITA containing acidic transactivator domain and NOD5 without any N-terminal domain.
1502:
44:
258:
There is also an additional subfamily NLRX which doesn't have significant homology to any N-terminal domain. A member of this subfamily is
1393:"Mutation of a new gene encoding a putative pyrin-like protein causes familial cold autoinflammatory syndrome and Muckle-Wells syndrome"
52:
1488:
1021:"NLRX1 is a mitochondrial NOD-like receptor that amplifies NF-kappaB and JNK pathways by inducing reactive oxygen species production"
1888:
1670:
143:
1560:
142:
Names as CATERPILLER, NOD, NALP, PAN, NACHT, PYPAF were used to describe the NLRs family. The nomenclature was unified by the
1256:"Influence of a nucleotide oligomerization domain 1 (NOD1) polymorphism and NOD2 mutant alleles on Crohn's disease phenotype"
160:
131:
1858:
1615:
1575:
1537:
1529:
56:
1898:
1663:
437:. These sugar chains are cross-linked by peptide chains that can be sensed by NODs. NOD1 recognizes a molecule called
98:. NLRs are highly conserved through evolution. Their homologs have been discovered in many different animal species (
397:
which leads to activation of RIP2. RIP2 mediates the recruitment of kinase TAK1 which phosphorylates and activates
469:
527:
1129:
Chen G, Shaw MH, Kim YG, Nuñez G (2009). "NOD-like receptors: role in innate immunity and inflammatory disease".
577:
118:(NOD or NBD – nucleotide-binding domain) domain, which is common to all NLRs, most of NLRs have also C-terminal
560:
463:
1555:
601:
523:
265:
On the other hand, NLRs can be divided into 3 subfamilies with regard to their phylogenetic relationships:
1903:
632:
626:
620:
453:
442:
79:
75:
1893:
576:
which indicate stress in the cell. The danger molecule can be extracellular ATP, extracellular glucose,
548:
406:
1494:
1525:
515:
434:
60:
1588:
1565:
655:
447:
438:
430:
119:
1545:
645:
606:
539:
509:
via PYD-PYD interaction. ASC contains PYD and CARD domain and links the NLRs to inactive form of
64:
43:) (also known as nucleotide-binding leucine-rich repeat receptors), are intracellular sensors of
410:
1484:
1463:
1422:
1373:
1332:
1287:
1236:
1187:
1146:
1102:
1050:
1001:
949:
900:
856:
815:
759:
708:
611:
458:
401:. The activation of IκB kinase results in the phosphorylation of inhibitor IκB which releases
1254:
Cantó E, Ricart E, Busquets D, Monfort D, García-Planella E, González D, et al. (2007).
1550:
1453:
1412:
1404:
1363:
1322:
1314:
1277:
1267:
1226:
1218:
1177:
1164:
Park JH, Kim YG, McDonald C, Kanneganti TD, Hasegawa M, Body-Malapel M, et al. (2007).
1138:
1092:
1040:
1032:
991:
983:
939:
931:
890:
846:
805:
797:
749:
739:
698:
688:
554:
1142:
835:"Nod2, a Nod1/Apaf-1 family member that is restricted to monocytes and activates NF-kappaB"
538:
There are three well-characterized inflammasomes – NLRP1, NLRP3 and IPAF. The formation of
589:
1019:
Tattoli I, Carneiro LA, Jéhanno M, Magalhaes JG, Shu Y, Philpott DJ, et al. (2008).
1872:
1417:
1392:
1282:
1255:
1231:
1206:
1166:"RICK/RIP2 mediates innate immune responses induced through Nod1 and Nod2 but not TLRs"
1045:
1020:
996:
971:
944:
919:
810:
785:
754:
727:
703:
676:
600:
irradiation and skin irritants. The presence of these molecules causes a production of
519:
91:
1882:
1639:
1207:"A critical role of RICK/RIP2 polyubiquitination in Nod-induced NF-kappaB activation"
801:
728:"Sweeten PAMPs: Role of Sugar Complexed PAMPs in Innate Immunity and Vaccine Biology"
677:"Sweeten PAMPs: Role of Sugar Complexed PAMPs in Innate Immunity and Vaccine Biology"
426:
414:
398:
149:
This system divides NLRs into 4 subfamilies based on the type of N-terminal domain:
1736:
1205:
Hasegawa M, Fujimoto Y, Lucas PC, Nakano H, Fukase K, Núñez G, et al. (2008).
970:
Ting JP, Lovering RC, Alnemri ES, Bertin J, Boss JM, Davis BK, et al. (2008).
650:
565:
505:
leads to the oligomerization. The pyrin domain of NLRs binds to an adaptor protein
490:
196:
127:
115:
48:
19:
987:
1829:
1695:
1643:
1182:
1165:
597:
585:
374:
170:
164:
123:
1867:
1097:
1080:
935:
95:
87:
83:
1272:
1222:
1036:
744:
693:
1820:
1623:
615:
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510:
68:
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1336:
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1240:
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1150:
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860:
851:
834:
819:
763:
712:
1350:
Srinivasula SM, Poyet JL, Razmara M, Datta P, Zhang Z, Alnemri ES (2002).
786:"Function of Nod-like receptors in microbial recognition and host defense"
593:
402:
1458:
1441:
1327:
1318:
920:"NOD-like receptors (NLRs): bona fide intracellular microbial sensors"
1810:
1805:
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1795:
1790:
506:
360:
356:
352:
348:
344:
252:
248:
244:
240:
236:
103:
1442:"[Biological and clinical aspects of Muckle-Wells syndrome]"
1391:
Hoffman HM, Mueller JL, Broide DH, Wanderer AA, Kolodner RD (2001).
879:"Human Nod1 confers responsiveness to bacterial lipopolysaccharides"
1352:"The PYRIN-CARD protein ASC is an activating adaptor for caspase-1"
522:. NLRP3 mutations are responsible for the autoinflammatory disease
1845:
1785:
1780:
1775:
1770:
1765:
1760:
1755:
1750:
1745:
1726:
1721:
1716:
1593:
1408:
833:
Ogura Y, Inohara N, Benito A, Chen FF, Yamaoka S, Nunez G (2001).
573:
543:
502:
498:
494:
405:
and its nuclear translocation. NF-κB then activates expression of
394:
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18:
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NLRPs and IPAF subfamilies are involved in the formation of the
274:
270:
178:
174:
24:
1659:
1498:
726:
Mahla RS, Reddy MC, Prasad DV, Kumar H (September 2013).
55:
that are associated with cell stress. They are types of
33:
nucleotide-binding oligomerization domain-like receptors
1856:
877:
Inohara N, Ogura Y, Chen FF, Muto A, Nuñez G (2001).
546:
such as microbial toxins (for example alpha-toxin of
1838:
1819:
1735:
1694:
1632:
1614:
1574:
1536:
604:and K+ efflux. NLRP1 recognizes lethal toxin from
1124:
1122:
1120:
1118:
1116:
1074:
1072:
1070:
1068:
1066:
1064:
972:"The NLR gene family: a standard nomenclature"
965:
963:
872:
870:
784:Franchi L, Warner N, Viani K, Nuñez G (2009).
779:
777:
775:
773:
45:pathogen-associated molecular patterns (PAMPs)
1671:
1510:
429:motifs from bacterial cell which consists of
94:and also in non-immune cells, for example in
8:
153:NLRA (A for acidic transactivating domain):
53:damage-associated molecular patterns (DAMPs)
918:Shaw MH, Reimer T, Kim YG, Nuñez G (2008).
1678:
1664:
1656:
1517:
1503:
1495:
580:, calcium pyrophosphate dihydrate (CPPD),
493:. The best characterized inflammasome is
59:, and play key roles in the regulation of
1457:
1416:
1367:
1326:
1281:
1271:
1230:
1181:
1096:
1044:
995:
943:
894:
850:
809:
753:
743:
702:
692:
457:). NOD2 proteins can sense intracellular
409:. Mutations in NOD2 are associated with
130:(PYD), acidic transactivating domain or
1863:
667:
524:familial cold autoinflammatory syndrome
1143:10.1146/annurev.pathol.4.110807.092239
23:Structure and domain organization of
7:
461:(MDP), typical for bacteria such as
57:pattern recognition receptors (PRRs)
552:) or whole pathogens, for instance
78:, whereas TLRs primarily recognize
14:
1446:Nihon Rinsho Meneki Gakkai Kaishi
114:NLRs contain 3 domains – central
102:) and also in the plant kingdom (
1866:
802:10.1111/j.1600-065X.2008.00734.x
144:HUGO Gene Nomenclature Committee
578:monosodium urate (MSU) crystals
1079:Schroder K, Tschopp J (2010).
67:and regulate inflammatory and
1:
1530:Pattern recognition receptors
588:or environmental irritants –
132:baculovirus inhibitor repeats
988:10.1016/j.immuni.2008.02.005
104:disease-resistance R protein
1183:10.4049/jimmunol.178.4.2380
441:(meso-DAP) mostly found in
307:NLRPs (also called NALPs):
27:, a human NOD-like receptor
1920:
1440:Kubota T, Koike R (2007).
1098:10.1016/j.cell.2010.01.040
477:Subfamilies NLRPs and IPAF
470:Mycobacterium tuberculosis
124:caspase recruitment domain
65:toll-like receptors (TLRs)
63:. NLRs can cooperate with
936:10.1016/j.coi.2008.06.001
497:, the activation through
74:NLRs primarily recognize
1273:10.3748/wjg.v13.i41.5446
1223:10.1038/sj.emboj.7601962
1037:10.1038/sj.embor.7401161
745:10.3389/fimmu.2013.00248
694:10.3389/fimmu.2013.00248
561:Saccharomyces cerevisiae
464:Streptococcus pneumoniae
439:meso-diaminopimelic acid
425:NOD1 and NOD2 recognize
47:that enter the cell via
1889:Intracellular receptors
1556:Formyl peptide receptor
732:Frontiers in Immunology
675:Mahla, Ranjeet (2013).
572:. NLRP3 recognize also
138:Nomenclature and system
1369:10.1074/jbc.C200179200
896:10.1074/jbc.M009728200
852:10.1074/jbc.M008072200
633:Listeria monocytogenes
627:Pseudomonas aeruginosa
621:Salmonella typhimurium
454:Pseudomonas aeruginosa
443:Gram-negative bacteria
407:inflammatory cytokines
80:Gram-negative bacteria
76:Gram-positive bacteria
61:innate immune response
28:
1260:World J Gastroenterol
549:Staphylococcus aureus
528:Muckle–Wells syndrome
22:
1526:Innate immune system
542:can be activated by
435:N-acetylmuramic acid
82:. They are found in
1589:RIG-I-like receptor
1566:Complement receptor
1538:Membrane-bound_PRRs
1459:10.2177/jsci.30.114
1081:"The inflammasomes"
656:RIG-I-like receptor
448:Helicobacter pylori
431:N-acetylglucosamine
120:leucine-rich repeat
1899:NOD-like receptors
1823:inhibitory protein
1561:Scavenger receptor
1546:Toll-like receptor
646:Toll-like receptor
607:Bacillus anthracis
540:NLRP3 inflammasome
37:NOD-like receptors
29:
1854:
1853:
1687:NOD-like receptor
1653:
1652:
1584:NOD-like receptor
1319:10.1002/art.24134
924:Curr Opin Immunol
612:muramyl dipeptide
459:muramyl dipeptide
16:Class of proteins
1911:
1871:
1870:
1862:
1839:Family member X1
1680:
1673:
1666:
1657:
1576:Cytoplasmic_PRRs
1551:Mannose receptor
1519:
1512:
1505:
1496:
1489:filter for human
1472:
1471:
1461:
1437:
1431:
1430:
1420:
1388:
1382:
1381:
1371:
1362:(24): 21119–22.
1347:
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1302:
1296:
1295:
1285:
1275:
1251:
1245:
1244:
1234:
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555:Candida albicans
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1633:Other/ungrouped
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1390:
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1348:
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1307:Arthritis Rheum
1304:
1303:
1299:
1266:(41): 5446–53.
1253:
1252:
1248:
1204:
1203:
1199:
1163:
1162:
1158:
1131:Annu Rev Pathol
1128:
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411:Crohn's disease
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92:dendritic cells
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1479:External links
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614:. IPAF senses
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381:Subfamily NODs
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51:or pores, and
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1091:(6): 821–32.
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545:
541:
533:
531:
529:
525:
521:
517:
512:
508:
504:
500:
496:
492:
485:Signalization
484:
482:
476:
474:
472:
471:
466:
465:
460:
456:
455:
450:
449:
445:(for example
444:
440:
436:
432:
428:
427:peptidoglycan
420:
418:
416:
415:Blau syndrome
412:
408:
404:
400:
396:
388:
386:
380:
376:
372:
368:
364:
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81:
77:
72:
70:
66:
62:
58:
54:
50:
46:
42:
38:
34:
26:
21:
1894:LRR proteins
1737:Pyrin domain
1686:
1583:
1449:
1445:
1435:
1403:(3): 301–5.
1400:
1396:
1386:
1359:
1355:
1345:
1310:
1306:
1300:
1263:
1259:
1249:
1214:
1210:
1200:
1173:
1169:
1159:
1134:
1130:
1088:
1084:
1028:
1024:
1014:
982:(3): 285–7.
979:
975:
927:
923:
913:
886:
882:
842:
838:
828:
793:
789:
735:
731:
721:
684:
680:
670:
651:Inflammasome
631:
625:
619:
605:
566:Sendai virus
559:
553:
547:
537:
507:ASC (PYCARD)
491:inflammasome
488:
480:
468:
462:
452:
446:
424:
392:
384:
264:
257:
195:NLRP (P for
169:NLRC (C for
159:NLRB (B for
148:
141:
128:pyrin domain
113:
73:
49:phagocytosis
40:
36:
32:
30:
1696:CARD domain
1644:Cyclophilin
1356:J Biol Chem
1328:2433/124253
883:J Biol Chem
839:J Biol Chem
790:Immunol Rev
586:cholesterol
395:kinase RIP2
88:macrophages
84:lymphocytes
71:response.
1883:Categories
1739:containing
1698:containing
1137:: 365–98.
662:References
399:IκB kinase
389:Signalling
96:epithelium
1821:Apoptosis
1624:Collectin
1485:PTHR14074
1397:Nat Genet
1170:J Immunol
616:flagellin
570:Influenza
511:caspase 1
110:Structure
69:apoptotic
1468:17473514
1427:11687797
1378:11967258
1337:19116920
1292:17907287
1241:18079694
1192:17277144
1151:18928408
1107:20303873
1055:18219313
1025:EMBO Rep
1006:18341998
976:Immunity
954:18585455
905:11058605
861:11087742
820:19120480
764:24032031
713:24032031
640:See also
594:asbestos
134:(BIRs).
126:(CARD),
1873:Biology
1418:4322000
1283:4171278
1232:2234345
1046:2267388
997:2630772
945:2572576
811:2679989
755:3759294
738:: 248.
704:3759294
687:: 248.
534:Ligands
421:Ligands
1859:Portal
1811:NLRP14
1806:NLRP13
1801:NLRP12
1796:NLRP11
1791:NLRP10
1713:/NLRC2
1707:/NLRC1
1689:family
1466:
1425:
1415:
1376:
1335:
1290:
1280:
1239:
1229:
1211:EMBO J
1190:
1149:
1105:
1053:
1043:
1004:
994:
952:
942:
903:
859:
818:
808:
762:
752:
711:
701:
590:silica
365:IPAF:
361:NLRP14
357:NLRP13
353:NLRP12
349:NLRP11
345:NLRP10
269:NODs:
253:NLRP14
249:NLRP13
245:NLRP12
241:NLRP11
237:NLRP10
1846:NLRX1
1786:NLRP9
1781:NLRP8
1776:NLRP7
1771:NLRP6
1766:NLRP5
1761:NLRP4
1756:NLRP3
1751:NLRP2
1746:NLRP1
1727:NLRC5
1722:NLRC4
1717:NLRC3
1594:RIG-I
618:from
574:DAMPs
544:PAMPs
520:IL-18
516:IL-1β
503:DAMPs
499:PAMPs
495:NLRP3
403:NF-κB
371:NLRC4
341:NLRP9
337:NLRP8
333:NLRP7
329:NLRP6
325:NLRP5
321:NLRP4
317:NLRP3
313:NLRP2
309:NLRP1
303:CIITA
299:NLRX1
291:NLRC5
283:NLRC3
260:NLRX1
233:NLRP9
229:NLRP8
225:NLRP7
221:NLRP6
217:NLRP5
213:NLRP4
209:NLRP3
205:NLRP2
201:NLRP1
191:NLRC5
187:NLRC4
183:NLRC3
155:CIITA
116:NACHT
100:APAF1
35:, or
1830:NAIP
1711:NOD2
1705:NOD1
1604:LGP2
1599:MDA5
1464:PMID
1423:PMID
1374:PMID
1333:PMID
1288:PMID
1237:PMID
1188:PMID
1147:PMID
1103:PMID
1085:Cell
1051:PMID
1002:PMID
950:PMID
901:PMID
857:PMID
816:PMID
760:PMID
709:PMID
610:and
582:alum
518:and
433:and
375:NAIP
367:IPAF
295:NOD5
287:NOD4
279:NOD3
275:NOD2
271:NOD1
179:NOD2
175:NOD1
171:CARD
165:NAIP
161:BIRs
41:NLRs
31:The
25:NOD2
1454:doi
1413:PMC
1405:doi
1364:doi
1360:277
1323:hdl
1315:doi
1278:PMC
1268:doi
1227:PMC
1219:doi
1178:doi
1174:178
1139:doi
1093:doi
1089:140
1041:PMC
1033:doi
992:PMC
984:doi
940:PMC
932:doi
891:doi
887:276
847:doi
843:276
806:PMC
798:doi
794:227
750:PMC
740:doi
699:PMC
689:doi
602:ROS
526:or
501:or
467:or
413:or
373:),
301:),
293:),
285:),
199:):
197:PYD
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