Theinnateimmunesystemworksasthefirstlineofdefenseinprotectionfrompathogenicmicrobesandhost-derivedsignalsofcellulardistress.Onewayinwhichthese“danger”signalstriggerinflammationisthroughactivationofinflammasomes,whicharemultiproteincomplexesthatassembleinthecytosolafterexposuretopathogen-associatedmolecularpatterns(PAMPs)ordanger-associatedmolecularpatterns(DAMPs)andresultintheactivationofcaspase-1andsubsequentcleavageofproinflammatorycytokinesIL-1betaandIL-18.Inflammasomecomplexestypicallyconsistofacytosolicpatternrecognitionreceptor(PRR;anucleotide-bindingdomainandleucine-rich-repeat[NLR]orAIM2-likereceptor[ALR]familymember),anadaptorprotein(ASC/TMS1),andpro-caspase-1.Anumberofdistinctinflammasomecomplexeshavebeenidentified,eachwithauniquePRRandactivationtriggers.ThebestcharacterizedistheNLRP3complex,whichcontainsNLRP3,ASC,andpro-caspase-1.TheNLRP3inflammasomeisactivatedinatwo-stepprocess.First,NF-kBsignalingisinducedthroughPAMP-orDAMP-mediatedactivationofTLR4orTNFR,resultinginincreasedexpressionofNLRP3,pro-IL-1beta,andpro-IL-18(primingstep,signal1).Next,indirectactivationofNLRP3occursbyamultitudeofsignals(wholepathogens,PAMPs/DAMPs,potassiumefflux,lysosomal-damagingenvironmentalfactors[uricacid,silica,alum]andendogenousfactors[amyloid-beta,cholesterolcrystals],andmitochondrialdamage),leADIngtocomplexassemblyandactivationofcaspase-1(signal2).Thecomplexinflammasomestructureisbuiltviadomaininteractionsamongtheproteincomponents.Otherinflammasomesareactivatedbymoredirectmeans:double-strandedDNAactivatestheAIM2complex,anthraxtoxinactivatesNLRP1,andbacterialflagellinactivatesNLRC4.Activatedcaspase-1inducessecretionofproinflammatorycytokinesIL-1betaand-18,butalsoregulatesmetabolicenzymeexpression,phagosomematuration,vasodilation,andpyroptosis,aninflammatoryprogrammedcelldeath.Inflammasomesignalingcontributestotheonsetofanumberofdiseases,includingatherosclerosis,typeIIdiabetes,Alzheimer’sdisease,andautoimmunedisorders.

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